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+The Project Gutenberg EBook of Ecological Observations on the Woodrat,
+Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
+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: Ecological Observations on the Woodrat, Neotoma floridana
+
+Author: Henry S. Fitch
+        Dennis G. Rainey
+
+Release Date: August 29, 2010 [EBook #33566]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK ECOLOGICAL OBSERVATIONS--WOODRAT ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci
+and the Online Distributed Proofreading Team at
+https://www.pgdp.net.
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+MUSEUM OF NATURAL HISTORY
+
+Volume 8, No. 9, pp. 499-533, 3 figs.
+
+June 12, 1956
+
+Ecological Observations on the Woodrat, Neotoma floridana
+
+BY
+
+HENRY S. FITCH AND DENNIS G. RAINEY
+
+UNIVERSITY OF KANSAS
+LAWRENCE
+1956
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson
+
+Volume 8, No. 9, pp. 499-533
+Published June 12, 1956
+
+UNIVERSITY OF KANSAS
+Lawrence, Kansas
+
+PRINTED BY
+FERD VOILAND, JR., STATE PRINTER
+TOPEKA, KANSAS
+1956
+
+
+
+
+ECOLOGICAL OBSERVATIONS ON THE WOODRAT, NEOTOMA FLORIDANA
+
+By
+
+Henry S. Fitch and Dennis G. Rainey
+
+
+_Introduction_
+
+The eastern woodrat exerts important effects on its community associates
+by its use of the vegetation for food, by providing shelter in its stick
+houses for many other small animals, and by providing a food supply for
+certain flesh-eaters. In the course of our observations on this rodent
+on the University of Kansas Natural History Reservation, extending over
+an eight-year period, from February, 1948, to February, 1956, these
+effects have changed greatly as the population of woodrats has
+constantly changed in density, and in extent of the area occupied.
+
+This report is concerned with the population of woodrats on the
+Reservation, the changes that the species has undergone, and the factors
+that have affected it. Our two sets of field data, used as a basis for
+this report, supplement each other and overlap little, either in time or
+space. Fitch's field work which covered approximately the western half
+of the Reservation, was begun in September, 1948, and was pursued most
+intensively in the autumn of 1948 and in 1949, with relatively small
+amounts of data obtained in 1950 and 1951 because of the great reduction
+in numbers of rats. Rainey's field work began in the spring of 1951 and
+was continued through 1954, concentrating on a colony in the extreme
+northwestern corner of the Reservation and on adjacent privately owned
+land. In actual numbers of rats live-trapped and for total number of
+records the two sets of data are comparable. Fitch's field work
+consisted chiefly of live-trapping while Rainey's relied also upon
+various other approaches to the woodrat's ecology. Rainey's findings
+were incorporated originally in a more comprehensive report (1956), from
+which short passages have been extracted that are most pertinent to the
+present discussion. Our combined data represent 258 woodrats (153
+Fitch's and 105 Rainey's) caught a total of 1110 times (660 Rainey's and
+450 Fitch's). Rainey's records pertain, in part, to woodrats outside the
+Reservation but within a few hundred yards, at most, of its boundaries.
+
+
+_Habitat_
+
+In the autumn of 1948 the population of woodrats was far below the level
+it had attained in 1947 or earlier, but the rats were still abundant and
+distributed throughout a variety of habitats. Almost every part of the
+woodland was occupied by at least a sparse population. Also, many rats
+lived beyond the limits of the woodland proper, in such places as
+deserted buildings, thickets, roadside hedges, and tangles of exposed
+tree roots along cut banks of gullies. All these situations are
+characterized by providing abundant cover, a limiting factor for this
+woodrat.
+
+In 1947, when the population of woodrats was especially high, plant
+succession on the wooded parts of the Reservation may have been near the
+optimum stage for the rats. For some 80 years, since the time the land
+was first settled and prairie fires were brought under control, woody
+vegetation has been encroaching into areas that were formerly grassland.
+
+About 1934 the University changed its policy with regard to treatment of
+the tract that was later made the Reservation. Up to that time, most of
+the area had been used as pasture and subjected to heavy grazing, but
+several fields had been fenced and cultivated. Under the new policy the
+hillsides and hilltop edges with open stands of various deciduous trees
+were enclosed with stock fences and protected from grazing. Successional
+trends were greatly altered. Woody vegetation, already favored by
+protection from the prairie fires originally important in the ecology of
+this region underwent further development as a result of protection from
+browsing. Thickets of shrubs and saplings sprang up throughout the
+woodland, forming a dense understory layer beneath the discontinuous
+canopy of the relatively scattered mature trees. The composition and
+density of the undergrowth varied markedly in different parts of the
+woodland. The parts that were formerly most open acquired the most dense
+understory. Blackberry, honey locust, osage orange, and prickly ash
+formed in places thorny tangles almost impenetrable to humans. This
+thicket stage reached its peak in density in the middle to late forties
+coinciding approximately with the time of maximum abundance of the rats.
+In the past eight years, under continued protection from burning,
+cutting and browsing, the forest has developed further; sizable trees 20
+feet or more high and up to eight inches in trunk diameter have grown
+from seedlings during the period of protection. An almost continuous
+canopy of foliage has developed, shading the understory and thinning it
+by killing shrubs and saplings. In those situations where the canopy is
+most dense, as on north slopes having stands of young hickory averaging
+twenty feet high, the understory is now largely lacking, but in other
+situations, particularly on south slopes, the understory thickets are
+still dense. On the whole, however, habitat conditions have become less
+favorable for the woodrat.
+
+Within the woodland the population of woodrats was not evenly
+distributed even at its maximum density; only those situations that
+provided sufficient overhead shelter were occupied by woodrats. The
+hilltop limestone outcrop, which was the refugium of the survivors when
+the population was at low ebb, also supported the greatest concentration
+when the population was high. The number of individuals living along any
+particular stretch of ledge could be determined only by intensive
+live-trapping, whereas residences of individuals could be more readily
+identified in most other situations away from the ledge. Stick houses of
+woodrats are, characteristically, large and dome-shaped in woodland, but
+along the ledges they usually lacked this typical form and consisted of
+a much smaller accumulation of sticks, often merely filling a small
+crevice. Sticks carried into such places where they were partly or
+wholly protected from moisture and sunshine were much less subject to
+decay than those in more open situations, and remained long after the
+rats themselves were gone. Accumulations of droppings in depressions in
+rock surfaces beneath overhanging ledges likewise have lasted for many
+years. The rock outcrop provided a continuous travelway along the
+hilltops, and even parts that were not permanently occupied usually had
+some sign. The following types of situations were found to be especially
+favorable for occupancy: deep crevices beneath overhanging projections
+of the ledge; large flat boulders broken away from the main ledge; thick
+clumps of brush (usually fragrant sumac, _Rhus trilobata_) providing
+shelter and support for the house; logs fallen across the ledge
+providing support and protection for the house structure.
+
+A second outcropping limestone stratum approximately 20 feet below the
+level of the hilltop was just as extensive as the upper outcrop, but it
+was little used by the rats because the exposed rock surface was more
+regular, lacking the jagged cracks and deep fissures of the hilltop
+outcrop; and it lacked the overhanging projections which provided
+overhead shelter for the rats along the upper outcrop. More than ninety
+per cent of the rats that were recorded as associated with the outcrops
+were at the hilltop stratum.
+
+Second in preference to the hilltop outcrop as a house site was the base
+of an osage orange tree in thick woods. This tree occurs throughout the
+woodland of the Reservation, having become established when the leaf
+canopy was more open, and the whole area was subject to grazing, with
+less development of the understory vegetation in the woodland. Houses
+were most often situated in those osage orange trees that had been cut
+one or more times, and had regenerated with spreading growth form, the
+multiple branching stems offering substantial support. Occasionally
+houses were built in crotches from two to six feet above ground.
+
+Blackberry thickets also are favorable locations for houses. These
+thickets grew up mostly in fenced areas from which livestock were
+excluded, but where there was not dense shade--hilltop edges and level
+or gently sloping ground adjacent to creek banks. The houses were
+usually in densest parts of the thickets where they were almost
+inaccessible. Mats of dead canes more or less horizontal, with the live
+canes growing up through them, provided effective overhead protection,
+while the ground beneath was relatively open. Houses built in the
+thickets were so well concealed that they were usually not detected
+until after leaves were shed in autumn. In most cases the blackberry
+thickets were small and well isolated. Houses of the rats were sometimes
+unusually near together suggesting that these thickets provided
+especially favorable habitat conditions.
+
+Hollow trees are often utilized, the accumulation of sticks for the
+house being largely inside the cavity. To be suitable for a house site,
+the snag must have an opening near ground level, and another higher on
+the trunk, providing emergency outlets in two directions. Most of the
+hollow trees utilized were black oaks (_Quercus velutina_).
+
+In 1948 there were many houses in cut tops of trees left from small
+scale lumbering operations a few years earlier. The densely branched
+tops of elms, oaks and hickories had satisfied the requirement for
+support of the house and nearby shelter. The houses built in them were
+in open woodland well separated from otherwise favorable situations. By
+1948 the tops were disintegrating and no longer provided effective
+shelter. The houses built in them were falling into disrepair and were
+not permanently inhabited but were often used temporarily by wandering
+individuals.
+
+Along cut banks of gullies where trees were partly undermined by
+erosion, the exposed, tangled root systems provided sites for occupancy.
+In these situations the accumulations of sticks were small and lacked
+the typical domed shape, consisting essentially of a lining to the
+cavity beneath the roots.
+
+Two small buildings at the Reservation headquarters were accessible to
+woodrats and were utilized off and on throughout much of the period of
+this study, despite the fact that most other sites of occupation away
+from the hilltop outcrops were deserted in the same period. One small
+building used as a laboratory had an enclosed wooden box five feet
+square housing an electric water pump. The interior of this box was
+accessible to the rats from beneath the floor. Litter of sticks and
+stems and various food materials were carried in by the rats. The nest
+thus protected and enclosed was not surrounded by the usual accumulation
+of sticks. An old garage 30 feet from the laboratory building was also
+occupied, sometimes by a different individual. The nest and food stores
+were behind boards propped against the wall.
+
+In October, 1948, live-trapping was begun on a heavily wooded slope
+facing northwest, and a ten-acre area was trapped rather thoroughly in
+the succeeding weeks. Because few traps were then available, this was
+the only area that was well sampled in 1948, although diffuse trapping
+was carried on over some 200 acres. On the ten-acre tract a total of 17
+adult and subadult woodrats were caught, four along the hilltop rock
+outcrop, six along the gully at the bottom of the slope, and seven at
+intermediate levels on the slope. Judging from the many unoccupied
+houses, the population on this tract had been much higher before the
+study was begun. On the basis of this sample it seems that in 1947 a
+population of several hundred woodrats lived on the wooded parts of the
+square mile where the Reservation is located.
+
+
+_Reduction of Population_
+
+The abrupt reduction in the population of woodrats on the Reservation
+cannot be explained conclusively with available data. Probably weather
+played a major part, but other unknown factors must have been important
+also. It is certain that the population of woodrats was high, if not at
+an all-time peak, in 1947. In late February, 1948, when one of us
+(Fitch) first visited the area on a preliminary inspection trip (not
+concerned primarily with woodrats), houses of these rats were found to
+be unusually numerous and those seen seemed to be occupied and well
+repaired. Possibly the population was drastically reduced within the
+next few weeks, as unseasonably cold and stormy weather occurred in
+early March. For the first 12 days of March, 1948, temperature averaged
+20° below that of average March weather, and even colder than the
+average for January or February. A reading of -5°F. on March 11 set a
+new low locally for the month since records were begun in 1869. The
+record low temperatures were accompanied by 12.8 inches of snow. This
+spell of unusually severe weather in early March coincided with the
+period in which first litters of young usually are born, as most females
+breed in early February and the gestation period is in the neighborhood
+of five weeks. That most of these first-litter young may have been
+eliminated by the unfavorable extreme of weather at the most critical
+stage in the life cycle may be readily imagined although definite proof
+is lacking. However, the mortality must have extended beyond newborn
+young. Loss of first litters ordinarily would be compensated for by the
+end of the season, since a female usually breeds more than once in the
+course of a season. In any case, by autumn, when the actual field study
+of woodrats was initiated, many houses were already deserted and in
+disrepair. Although the rats were still moderately abundant, they were,
+seemingly, much below the population peak of the preceding year.
+
+Further drastic reduction of adults and subadults took place in the
+winter of 1948-49. In the course of live-trapping operations from
+mid-October into early December, 51 individuals were caught and marked.
+Chiefly because of unfavorable weather conditions, field work was
+discontinued in mid-December, and live-trapping was not resumed until
+early March. Subsequently, only 12 of the woodrats previously marked
+could be recaptured, and the population had become noticeably sparse.
+Seemingly, more than three-fourths of the population present in late
+autumn had been eliminated in the interval. In January, weather was
+exceptionally severe; on the ninth and tenth the worst sleet storm in
+twelve years occurred. Sleet fell in small granules, while the
+temperature remained several degrees below freezing. Partial thawing on
+January 12, 13 and 14 was followed by a steady drizzling rain on the
+fifteenth. On the following day the temperature dropped to -7°F. Ice
+still remained from the sleet storm, and the slush again froze. On the
+night of January 18, there was one of the worst snow storms on record
+and temperature reached a low of 2°F. Exceptionally low temperatures
+persisted through January 24, with more sleet on January 25. Ice from
+the earlier storm still remained. On January 30, the temperature dropped
+to -7° and a three-inch cover of snow still remained over the coat of
+ice. The month of January, 1949, had the heaviest precipitation in 81
+years (5.09 inches) and a cover of ice remained for at least 21 days.
+There were other sleet storms of lesser proportions on February 2 and
+again on February 21.
+
+Ordinarily sleet would not seriously damage woodrats living in houses in
+woodland habitats and less suitable hedge rows because it usually
+freezes as it falls and coats only the surface of the house. Gradual
+thawing would allow normal runoff without much penetration. Because the
+sleet during the storm described above did not form a glaze as it fell,
+the ice particles penetrated many houses. It has been observed many
+times that captive woodrats refused food that was frozen or were unable
+to eat it. Woodrats in live-traps in winter rapidly weaken unless a
+large supply of food is available. If food supplies became sealed over
+by ice, woodrats would have died by starvation or by falling an easy
+prey to predators. The rats were more accessible to several predators
+than were smaller mammals such as meadow voles which were difficult to
+obtain because of the coating of ice over the fields.
+
+The decimated population surviving into the breeding season of 1949
+failed to make substantial gains. In fact, during the following
+four-year period the general trend of the population over the
+Reservation as a whole seemed to be one of gradual further decline.
+
+In November, 1949, the rats were almost gone from the area of north
+slope and hilltop in oak-hickory-elm woodland where the most intensive
+live-trapping and other field work had been done the previous year. The
+following descriptions of houses remaining on the area at that time give
+some idea of the habitat, and of the course of events correlated with
+the fluctuations in numbers of woodrats.
+
+     No. 1. At the hilltop outcrop, partly on a substrate of
+     limestone boulders, built around an elm of two-foot DBH,
+     which lent support to one side. A hackberry sapling one inch
+     in stem diameter grew through the middle of the house,
+     providing further support. The house was two feet high and
+     six feet in diameter, and was in obvious disrepair, with a
+     hole several inches in diameter in its top. It had been
+     occupied in the autumn of 1948. It was constructed mainly of
+     sticks, ranging in diameter from approximately one inch to
+     straw size. Many of the sticks, from .4 to .5 inches in
+     diameter and one to two feet long, seemingly would have been
+     heavy burdens for a rat, although they were of light-weight
+     wood, sumac and elm. Mixed with the sticks were quantities
+     of dry leaves, bark, and chips of wood, all material
+     appearing old and weathered. This house was in
+     elm-oak-hickory woods 50 feet from a cultivated field on the
+     hilltop to the east and south. To the north and west the
+     escarpment sloped away abruptly. There was a coralberry
+     thicket beneath the trees on the adjacent hilltop.
+
+[Illustration]
+
+FIGURE 1
+
+(A) Map of part of University of Kansas Natural History Reservation,
+showing first-capture sites for all woodrats live-trapped in the autumn
+of 1948. Because of the short time involved and the few traps available,
+much of the area shown was not thoroughly trapped. Woodrats were
+abundant, though much less so than in 1947, as shown by the large number
+of deserted houses.
+
+(B) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1949. Woodrats were
+still moderately abundant, but much below the level of the previous
+year. Triangles indicate those capture sites not sampled in 1948.
+
+(C) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1950. Numbers were
+medium-low, having undergone drastic reduction from the peak level.
+Triangles indicate those capture sites where trapping was not done in
+earlier years.
+
+(D) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1951. The
+population was low, but had not yet reached its lowest ebb.
+
+(E) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1952, when the
+population had declined to relatively low numbers and disappeared from
+much of its former habitat.
+
+(F) Map of the 590-acre Natural History Reservation, showing the area
+where woodrats were studied.
+
+     No. 2. On gently sloping hilltop edge 15 feet from the
+     outcrop and escarpment, built around a forked walnut sapling
+     having both trunks approximately five inches in diameter.
+     The sapling, coming up through the center of the house at a
+     45° angle, evidently had been bent by the accumulated weight
+     of the debris at an early stage of its growth, many years
+     before. Trees were small in this part of the woods, with a
+     well developed understory thicket of coralberry and sumac.
+     This house approximately one foot high and six feet wide,
+     was constructed mainly of sticks and was similar in
+     composition to No. 1, but appeared considerably older with
+     all the sticks blackened and rotten. In the autumn of 1948
+     this house was used by woodrats, but probably only as a
+     temporary stopping place, because it was already in
+     disrepair then.
+
+     No. 3. At edge of escarpment, 25 feet from No. 2, on a flat
+     boulder approximately six feet long, three feet wide and one
+     foot thick. The decaying and much flattened mass of sticks
+     was mainly on top of the boulder, but also spilled over its
+     edges. Fresh sign was noted at this house in the autumn of
+     1948, but the house was already in disrepair then, and
+     seemingly it was used only as a stopping place.
+
+     No. 4. At the hilltop outcrop where an elm had fallen across
+     it. The decaying log remaining was approximately 12 feet
+     long and 15 inches thick. This log passed diagonally through
+     the house, providing its main support. The house was
+     approximately 39 inches high, its summit extending a little
+     above the level of the top of the outcrop. The house was
+     approximately seven feet wide along the outcrop. This house
+     was somewhat intermediate between the typical dome-shaped
+     stick piles that the rat builds in open situations and the
+     formless accumulations of sticks with which some rats living
+     in deep rock crevices line the entrances. Part of the
+     accumulation was beneath the limestone boulders and
+     outcropping slabs. Approximately half of the material used
+     in the house consisted of sticks and the remainder of pieces
+     of bark and chips of wood, mostly gathered from the fallen
+     elm. This house had shrunken noticeably from decay and
+     settling in the months since it was occupied, in the autumn
+     of 1948. The house was surrounded by a thicket of fragrant
+     sumac, dogwood, and hackberry saplings.
+
+     No. 5. At edge of a protruding boulder one foot thick at the
+     hilltop outcrop of the west facing escarpment, and 100 feet
+     back in the woods from the edge of a corn field, in
+     undergrowth of dogwood, wild currant, and coralberry. The
+     house consisted of a pile of rotten twigs, 3 inches deep and
+     30 inches wide on the upper side of the boulder, and a
+     lining of similar material at the lower edge of the boulder,
+     partly blocking the crevice beneath it. The twigs composing
+     the house were old and rotten. However, a few dry but still
+     green hackberry leaves were stored in the crevice beneath
+     the boulder. In a bare space atop the boulder were several
+     recent woodrat droppings, small and obviously produced by an
+     immature individual, which, perhaps, had recently settled at
+     this old house site.
+
+     No. 6. In hilltop woods, 30 feet from a corner adjoining a
+     pasture and a corn field, at the base of an osage orange
+     tree of one foot DBH, and also over a hollow cottonwood log
+     one foot in diameter and three feet from the osage orange
+     tree. Suspended mats of grape and smilax vines, and the
+     thorny, dead, lower branches of the tree provided additional
+     shelter. The house was composed of sticks and twigs, mostly
+     of osage orange, with spines still present; slabs of bark,
+     wood chips, and dry leaves also made up part of it.
+     Materials on the exterior of the house appeared old and
+     weathered, but the house was conical and solid. Seven fresh
+     corn cobs were on the house or near its base, suggesting
+     that corn from the nearby field had figured importantly in
+     the diet of the occupant. A well beaten path led from the
+     base of the house alongside the log, to a large cottonwood
+     tree 15 feet from the house. This evidence that the house
+     was occupied was verified by live-trapping the occupant.
+     Late in 1948, also, the house was occupied by another
+     individual, but seemingly was deserted for a period of
+     months thereafter.
+
+     No. 7. On upper part of north slope where a hickory seven
+     inches in diameter had fallen across an old sunken log
+     approximately one foot in diameter. The house, composed
+     mainly of hickory twigs 1/4-inch to 1/2-inch in diameter,
+     mixed with bark, wood chips, and leaves, was partly decayed,
+     with no fresh sign and was in a thicket of greenbrier,
+     saplings of hickory and hackberry, and cut tops of
+     hickories. The top was flattened to less than four inches
+     above the level of the supporting hickory log. There were
+     large cavities in the side of the house. When first
+     discovered in the autumn of 1948, this house was occupied by
+     a subadult female rat, but she moved away permanently, and
+     the house had been deserted for approximately a year when
+     these observations were recorded.
+
+     No. 8. In middle of northwest slope, in thick branches of
+     broken top of a black oak. This house had become flattened
+     by decay and settling to form a mound approximately one foot
+     high and five feet in diameter. Only the top protruded
+     through the carpet of dry leaves. Once well protected and
+     partly concealed by the branches and twigs of the oak top,
+     this house was now fully exposed by the disintegration of
+     the top. The house consisted chiefly of oak twigs. In
+     October, 1948, a woodrat was live-trapped at this house, but
+     probably it was a wanderer. The house had then already
+     undergone much deterioration.
+
+
+_Natural Enemies_
+
+Some 56 species of animals that regularly prey on small vertebrates live
+on the Reservation. Many of the larger kinds may take woodrats
+occasionally. Because of size, habitat preferences and the time and
+manner of hunting, five species stand out as the more formidable
+enemies--the horned owl (_Bubo virginianus_), prairie spotted skunk
+(_Spilogale putorius_), long-tailed weasel (_Mustela frenata_), pilot
+black snake (_Elaphe obsoleta_) and timber rattlesnake (_Crotalus
+horridus_).
+
+Throughout the study horned owls were common on the area, but their
+numbers were highest in 1948. Samples of pellet collections have shown
+that the cottontail is the staple food, being represented in almost
+every pellet. Various rodents also are important in the diet, the cotton
+rat, prairie vole, or white-footed mouse being most prominent according
+to the time and place of collection. The woodrat is approximately
+optimum size for prey, and it constitutes one of the most preferred food
+sources. Remains of only two woodrats were found in the pellets
+examined, but at times when the pellets were collected woodrats were so
+scarce that they constituted only an insignificant percentage of the
+biomass of potential prey. On several occasions woodrats in live-traps
+were attacked by horned owls, as shown by the overturned and displaced
+trap and quantities of fine down adhering to them and to nearby objects.
+The horned owl lives in the same habitat as does the woodrat. In other
+regions woodrats are known to figure prominently in the diet of the
+horned owl. At the San Joaquin Experimental Range in California, for
+instance, _N. fuscipes_ was found 240 times, more frequently than any
+other kind of prey, in 654 pellets of the horned owl, and this owl was
+shown to be the one most important natural enemy of the rat, although
+many kinds of carnivores, raptors and snakes also took toll from its
+populations. On the Reservation the population of horned owls has been
+fairly stable from year to year, with roughly one pair to 100 acres of
+woodland. Some territories have been maintained continuously throughout
+the eight-year period of observation, though changing to some extent in
+size, shape and area included. In 1948, when livestock grazed on the
+area, and the ground cover of herbaceous vegetation was relatively
+sparse, cottontails were much less abundant than they were later when
+the vegetation was protected. Small rodents including voles, cottonrats,
+and deer mice, were also less abundant then, and the numerous horned
+owls may have been supported in part by the high population of woodrats.
+
+The spotted skunk may be an even more important enemy of the woodrat,
+although the evidence is circumstantial. No records of these skunks
+preying on woodrats have been found in the literature, nor were any such
+instances recorded by us except for attacks on woodrats confined in
+live-traps. This skunk is a formidable enemy of small and medium-sized
+rodents, as it can climb, dig, and squeeze through small openings. That
+it may prey on rat-sized rodents and may even be a limiting factor to
+their occurrence is well shown by Crabb's (1941:353) studies in Iowa. He
+found that Norway rats (_Rattus norvegicus_) ranked third in frequency
+(cottontail, mostly carrion, ranked first) in the winter food of the
+spotted skunk. Crabb observed that about farmyards and farm buildings
+where the skunks had been eliminated by persistent persecution, rats
+were abundant, but that about others where the skunks were present, the
+rats were scarce or absent. On several occasions he noted that heavy
+populations of rats about farm buildings in summer and autumn nearly
+disappeared in winter if a skunk was in residence.
+
+Sign of spotted skunk was noted frequently on various parts of the
+Reservation, especially along the hilltop ledges which were the best
+woodrat habitat. On several occasions skunks released from live-traps
+took shelter in woodrat houses which appeared to be unoccupied.
+According to a local fur dealer, C. W. Ogle, spotted skunks reached a
+peak of abundance in Douglas County in the winter of 1947-1948, and many
+pelts were brought in for sale then. The concentration of skunks may
+have had detrimental effect on the population of woodrats, especially
+when extremes of weather had already made conditions critical for them,
+as in early March, 1948, and in January, 1949, when snow and sleet made
+their usual food supply unavailable.
+
+The long-tailed weasel is considered to be a potentially important enemy
+of the woodrat. Weasels have been seen on the Reservation on only a few
+occasions, but they may be more numerous than these records would
+indicate. Two were caught at the hilltop outcrop, at different times and
+places, in funnel traps put out to catch snakes. The weasel seems to
+prefer this rocky habitat, which is also favored by the woodrat. Because
+of its ferocity and willingness to attack relatively large prey, and
+because it is an agile climber and able to squeeze through any openings
+large enough to accommodate a woodrat, it would seem to be a formidable
+enemy.
+
+The pilot black snake (_Elaphe obsoleta_) is an important enemy of this
+woodrat on the Reservation and probably throughout the rat's geographic
+range except for the extreme western part. Although this snake occurs in
+every habitat of the Reservation, it has been found most often along
+rock outcrops of wooded hilltop edges in the type of habitat most
+favored by the rat. Most often pilot black snakes have attempted to
+escape into crevices of the outcrop. These snakes are also skillful
+climbers and often have escaped by climbing out of reach along branches
+or even vertical tree trunks. On several occasions these snakes have
+been found on or beside woodrat houses, or have escaped into them. Over
+a seven-year period 143 pilot black snakes have been recorded, 53 of
+which were adults.
+
+On September, 1948, a large pilot black snake found basking on a rock
+ledge, distended by a recent meal, was palped and contained a subadult
+female woodrat. On June 19, 1953, one of us, approaching a live-trap set
+under an overhanging rock ledge, saw a four-foot pilot black snake on
+top of it. The snake struck repeatedly at the rat in this trap, but was
+unable to reach it. At each stroke the rat would dash about the trap
+frantically.
+
+These snakes hunt by stealth, and might catch woodrats by entering their
+nests, or by lying in wait along their runways, but are not quick enough
+to catch them in actual pursuit. Young in the nest would seem to be
+especially susceptible to predation by the pilot black snake. These
+snakes hunt by active prowling, either by night or by day, and much of
+their food consists of the helpless young of birds and mammals found in
+the nests. While only well-grown or adult pilot black snakes would be
+able to swallow an adult woodrat, any but first-year young probably
+would be able to overcome and swallow the small young. The female
+woodrat's habit of dragging the young attached to her teats as she flees
+from the house at any alarm must save many litters from predation by the
+pilot black snake. First litters of young, born in early March, are
+already well grown, and past the age of greatest susceptibility to
+predation before the snakes emerge from hibernation in late April or
+early May.
+
+The timber rattlesnake is another potentially destructive enemy, but on
+the Reservation, and throughout much of its original range it is now
+relatively scarce. The genus _Neotoma_ largely coincides in its over-all
+distribution with the genus _Crotalus_, of the rattlesnakes. For most
+kinds of woodrats, the larger species of rattlesnakes are among the
+chief natural enemies.
+
+The timber rattlesnake has habitat preferences similar to those of the
+eastern woodrat. Of 30 timber rattlesnakes recorded on the Reservation
+over an eight-year period, all but one were at or near hilltop rock
+ledges in woodland. The woodrat is probably one of the most important
+prey species for the timber rattlesnake. Like the woodrat, the
+rattlesnake is mostly nocturnal in its activity. Unlike the pilot black
+snake, it hunts by lying in wait, striking prey which comes within
+range, and waiting for it to die from the venomous bite, rather than by
+active prowling. Therefore, it is probably less of a hazard to young in
+the nest than is the pilot black snake. Even young rattlesnakes too
+small to eat woodrats are potentially dangerous to them, as they may
+strike and kill any that come within range.
+
+
+_Commensals_
+
+Rainey (1956) listed many kinds of small animals that use the houses of
+the eastern woodrat and live in more or less commensal relationships
+with these rodents.
+
+A situation unusually favorable for observing woodrats and their
+associates was discovered on the Reservation where, in July, 1948, two
+old strips of sheet metal, each covering an area of approximately 25
+square feet, were used as shelter by a lactating female with three
+young. This was on a brushy slope just below an old quarry site. A rock
+pile and remains of an old rock wall were nearby. Woodrats had carried
+many sticks back under the metal strips, filling the spaces beneath
+their edges. There was a nest and a system of runways beneath the
+strips. In the following seven years this site was seldom deserted for
+long and was used by a succession of individuals. The strips of metal
+could be easily raised and then lowered into place with little
+disturbance. Because the situation was not entirely natural, the
+findings may not be typical of other rat houses. Animals found over a
+period of years beneath these metal strips include: several dozen each
+of the ring-necked snake (_Diadophis punctatus_), five-lined skink
+(_Eumeces fasciatus_), and ant-eating toad (_Gastrophryne olivacea_);
+several individuals each of cottontail (_Sylvilagus floridanus_),
+white-footed mouse (_Peromyscus leucopus_), short-tailed shrew (_Blarina
+brevicauda_), least shrew (_Cryptotis parva_), American toad (_Bufo
+americanus_), Great Plains skink (_Eumeces obsoletus_), pilot black
+snake (_Elaphe obsoleta_); and one each of bull snake (_Pituophis
+catenifer_), spotted king snake (_Lampropeltis calligaster_), red milk
+snake (_L. triangulum_), and timber rattlesnake (_Crotalus horridus_).
+The snakes which were potential predators on the rats seemed to be
+merely utilizing the shelter in these instances, but they may have been
+lying in wait for prey there.
+
+Among mammals, the cottontail and the white-footed mouse are the most
+persistent users of the woodrat houses, especially those that are no
+longer occupied by the rats. On one occasion five white-footed mice were
+caught simultaneously in a trap set beside a house at the base of an
+osage orange tree. Subsequent trapping showed that this house was no
+longer occupied by a rat, but that the mice lived in it. Occupancy of
+such an old woodrat house by white-footed mice may continue long after
+abandonment of the house by the rat, even after the house has partly
+decayed and settled to a small part of its original volume.
+
+Cottontails often have their forms under the edges of houses, either
+occupied or deserted. These situations offer protection overhead and on
+three sides. Abandoned houses having one or more of the entrance holes
+enlarged, as by predators breaking through the side of the house to gain
+access to the nest, are especially well adapted for occupancy by the
+cottontail. The rabbit may make its form inside the house structure.
+
+The opossum, also, finds the type of shelter that it requires in
+abandoned houses that have had the entrances sufficiently enlarged. On
+various occasions opossums or their remains have been found in such old
+houses, and opossums released from live-traps have been known to seek
+shelter in abandoned woodrat houses.
+
+At the old quarry on the Reservation woodrat sign was especially
+abundant. A wooden bin approximately seven feet square, used to store
+crushed rock before quarrying operations were abandoned, was inhabited
+by one rat. At the base of a rock crusher on the top of a bank a few
+yards from the bin was an accumulation of sticks and other debris
+brought by woodrats. A rock wall at the top of the bank between the
+crusher and the bin had many crevices providing shelter for the rats,
+and projecting rocks were littered with their droppings. In the spring
+of 1949 the bin and rock crusher were removed, but at least one rat
+continued to live in the rock wall. In the summer of 1951 several tons
+of corn ruined in the flood were dumped on the top of the bank above the
+wall. By autumn, Norway rats, either brought in with the corn or
+attracted by it, had taken possession of the wall, evidently displacing
+the woodrats, which were no longer present. Although this Old World
+murid rat is much different from the woodrat in habits, it seemingly can
+compete with it and replace it where habitat conditions are otherwise
+favorable for both.
+
+
+_Movements_
+
+The woodrat is dependent on the stick houses that it constructs for
+shelter. For each individual the house constitutes a home base to which
+it is attached, and about which its movements revolve. The area within
+which routine daily movements are confined constitutes the home range,
+which is variable in size and shape. An individual may, and usually
+does, alter its home range over periods of time. The home range is
+somewhat nebulous because the rat may at any time move far beyond the
+small area to which its activities are largely confined. It may be
+motivated by sexual urge or other voluntary wandering; it may be
+enticed by a food supply or some other specific attraction not available
+near its house; or it may be forcibly displaced by an intruder or may
+abandon in favor of an offspring.
+
+An occupied house normally has several runways radiating from it. These
+are well worn paths, smoothed by use, and cleared of obstructions, and
+the rat tends to keep to them in its foraging expeditions. Usually a
+trail leads to a bush or tree showing evidence of heavy use by the rat.
+Ordinarily such a trail cannot be traced more than 30 feet from the
+house, and it seems that the most concentrated foraging occurs within
+this short radius. Experience in live-trapping has indicated that the
+distance covered by a woodrat in its normal foraging for food is
+ordinarily less than 75 feet in any direction from the house.
+
+Usually the rats can be caught in traps only at their houses or nearby
+places that they frequent, as indicated by their sign. When travelling,
+woodrats make use of overhead cover as much as possible. Storing of food
+seems to be associated with the animal's reluctance to wander far from
+home. When a rat is gathering preferred food for storage the home range
+may be enlarged (or the animal may travel beyond the limits of its
+regular home range). In any case the rat may find it necessary to
+traverse an additional area in order to reach the food source. This may
+involve, in part, extension vertically, as when the rat obtains food
+from trees directly over the house. The home range is thus somewhat
+three-dimensional; both trails and feeding places are often above
+ground. Because of dependency on cover, woodrats do not forage randomly
+in all directions from the house.
+
+Although the house and its immediate environs are defended as a
+territory by the occupant, possession may be soon relinquished. A
+woodrat may shift frequently from one house to another, especially if
+unoccupied houses are readily available. Because woodrats had undergone
+drastic reduction in numbers, as discussed on p. 505, unoccupied houses
+in various stages of disrepair were numerous throughout the woodland in
+1948 and 1949, and the rats that were present then seemed especially
+inclined to wander. Even old houses that are collapsed and
+disintegrating may be used temporarily, or may be taken over and
+repaired. Houses that are in sites exceptionally favorable in that they
+provide food and shelter may be occupied more or less permanently, with
+a succession of woodrats over many generations.
+
+Shifts to new areas are perhaps most often motivated by a search for
+mates. Such shifts are, on the average, longer and more frequent in
+males. Males must range farther in search of females when numbers are
+low. On the other hand, when numbers are high and most of the best sites
+are occupied, newly independent young and displaced adults are forced to
+travel greater distances in search of homes. Some of the larger and more
+powerful males move far greater distances than smaller males. The
+longest distances recorded were mostly for large adult males in breeding
+condition. The average maximum distance between successive points of
+capture for 27 adult males was 345 feet. For 39 females (adults and
+subadults) the corresponding figure was 143 feet. The extremes for males
+were 0 to 1080 feet and for females, 0 to 650 feet. Of the 27 males,
+five moved the maximum distance in a single night. Most of the long
+movements by males did not constitute clear-cut shifts in home range,
+and many returned to their original locations.
+
+The average distance between points of first and last captures for 72
+subadult and adult males was 165 feet. A similar figure for 72 subadult
+and adult females was 133 feet. Of the males 23.7 per cent were at the
+same place at the first and last captures; for females the percentage
+was 36.1. These figures are from the combined data of our trapping
+records, but the trends differed sharply in the two sets of records. In
+Fitch's records, movements averaged longer and difference between the
+sexes was much less: 189 feet for 41 males and 178 feet for 42 females.
+Corresponding figures from Rainey's records were: 141 feet for 31 males
+and 74 feet for 30 females. In Fitch's field work, opportunities to
+record exceptionally long movements obviously were better because the
+trap line encompassed a larger area, approximately half a square mile,
+whereas Rainey's live-trapping was concentrated on relatively small
+areas. The reason for the greater vagility of females in Fitch's records
+is less evident. However, the data were obtained within the period of
+drastic population reduction, at a time when there were numerous empty
+houses throughout the woodland, facilitating travel, and shifts from one
+home range to another where conditions were, temporarily at least, more
+favorable. Rainey found that the females in the small colony in woodland
+where he trapped, moved much less than did those that lived along the
+hilltop outcrop, which provided a natural travel route.
+
+Following are several examples of males and females with long histories
+showing individual variation in frequency and distance of movements.
+
+     _Males_
+
+     (1.) First captured October 14, 1951, and last captured 327
+     days later on September 6, 1952. He was taken 12 times. For
+     the first seven captures (October 14, 1951, to July 15,
+     1952), no movements were recorded. In the following seven
+     days he moved 367 feet. Within the next 21 days he returned
+     to within 114 feet of the site of original capture. Less
+     than one month later he was caught for the last time, at
+     this same site.
+
+     (2.) This large male was captured twelve times over a period
+     of 827 days (March 16, 1952, to June 21, 1954). He tended to
+     wander more than other males and was absent from the
+     trapping area from early 1952 to May 1953. One round trip
+     made in a two-weeks period, amounted to a linear distance of
+     1894 feet if the rat followed natural cover. The return trip
+     of 947 feet was the greatest distance traversed in a single
+     night in any of the woodrats we recorded. Other movements
+     between successive captures were: 722, 397, 356, 293, 253
+     and 144 feet (the latter shift made three different times).
+     Sexual urge probably motivated most of his wandering, since
+     numbers of females were low.
+
+     (3.) For this male the span of records was 143 days, with 18
+     captures. For the first eight recaptures, extending over a
+     period of 39 days, he was still at the original location.
+     Four days later he had moved 120 feet and was visiting a
+     female. A week later he returned. In the following month he
+     was recorded as making two more moves, of 115 feet and 215
+     feet. He was last recorded at the hilltop outcrop.
+
+     (4.) The records of this male extended over 465 days, with
+     13 captures. For the entire period only one movement, of 163
+     feet, was recorded. Twelve of the 13 captures were at the
+     same house.
+
+     (5.) This male was captured 16 times over a span of 130
+     days. After the second capture he moved 144 feet along the
+     outcrop and was caught there for the next 14 times, having
+     developed a "trap habit."
+
+     (6.) This male was in the area 210 days (13 captures) and
+     shifted his range. He was first captured on August 17, 1952,
+     at a house at the rock fence 433 feet from the outcrop.
+     Between this date and October 12, 1952, he moved to the
+     outcrop and established residence in a vacant house. He was
+     recorded as making six more moves, the longest of which was
+     only 40 feet.
+
+     (7.) This male was first caught in June, 1949, as a juvenile
+     probably between two and three months old (weighing 96
+     grams) and hence probably still at the maternal house. In
+     September, grown to adult size, he was caught twice, still
+     at this same place. In October, November, December, and in
+     February, 1950, he was caught 11 times at eight places all
+     within a 90-foot radius of his original location. In April,
+     1950, he was caught at points 550 feet WSW and 700 feet SW.
+     In October he was caught within 150 feet of the original
+     location. In November, 1950, and in March and April, 1951,
+     he was caught four times at a place 900 feet SW from his
+     original location.
+
+     (8.) This subadult male was first caught at the hilltop
+     outcrop on October 4, 1949. Two days later he had moved 160
+     feet north along the outcrop. A month later he had shifted
+     600 feet south; in three more days 1040 feet north. On
+     November 15 he was 105 feet south of the November 8
+     location; on November 16, he had moved 70 feet north. On
+     November 17 he had moved 900 feet back south, but had
+     returned on the 18th to the November 16 location. On
+     November 22, he had again shifted 900 feet south. All
+     capture sites were at the hilltop outcrop.
+
+     (9.) This male was caught as a juvenile (75 grams) on
+     October 8, 1950. On November 9 he had moved 220 feet, from
+     the lower outcrop to the upper, and he was recaptured at or
+     near this same site on November 10, 28 and 29, and on
+     January 11 and February 9, 1951. On November 21, 1951, grown
+     to maximum adult size, he was caught at a new location 1080
+     feet from the original.
+
+     (10.) This male was caught as a subadult twice at the same
+     place on November 30 and December 14. By the following
+     autumn he had shifted to a new location 180 feet south along
+     the outcrop, and he was caught there on September 22 and
+     October 18, 1951, and on January 20 and February 2, 1952.
+
+
+     _Females_
+
+     (11.) This female was captured 27 times over a span of 211
+     days. She moved back and forth considerably between two
+     houses 40 feet apart but made only one substantial movement
+     of 245 feet; at this time she was in breeding condition.
+     Nearly seven months after the first capture she was seen for
+     the last time only 16 feet from the original site of
+     capture. It was assumed she fell prey to spotted skunks
+     which were raiding traps.
+
+     (12.) First captured on March 24, 1951, she remained on the
+     area 105 days in which period she was live-trapped 25 times.
+     Sixty per cent of the total captures were at the same house
+     and the longest movement recorded was only 56 feet. She was
+     last caught in a trap 25 feet from the site of original
+     capture.
+
+     (13.) This young adult remained at her house at the rock
+     fence approximately four months. In this period she was
+     captured 11 times. On March 16, 1952, she had moved 410 feet
+     to a house at the eastern section of outcrop, probably
+     searching for a male. She was never seen again.
+
+     (14.) This subadult female moved from the site of original
+     capture to a house 253 feet away on the same outcrop. She
+     was probably in search of a new home when caught the first
+     time. She was recorded at another house 40 feet away on one
+     occasion.
+
+     (15.) Over a span of 90 days and 15 captures this female was
+     not recorded as making any movement. She was living in one
+     of the woodland houses. Mature males were numerous in the
+     area and she was visited by at least two.
+
+     (16.) This female was also living in the woodland section
+     and was first caught on March 30, 1952, in one of the less
+     favorable houses. She was trapped 17 times over a period of
+     85 days. One movement of 68 feet to a new home site was
+     recorded, but the area of foraging probably did not change.
+     She was caught here four times and then disappeared.
+
+     (17.) This female was first trapped as a subadult on October
+     5, 1948, at a house in brush on the upper part of a north
+     slope. On November 24 she had shifted 590 feet to the bottom
+     of the slope and was living in the recess beneath an
+     undermined honey locust on a gully bank. On November 25 she
+     was caught in a similar situation 100 feet farther east
+     along the gully bank. She was recaptured at the gully on
+     November 26 and 30, December 1, 3, 22, and March 8 and 9,
+     and in all she shifted six times between the two gully-bank
+     dens.
+
+     (18.) This female was first trapped as an adult on November
+     18, 1948, in a gully-bank den. She was recaptured at this
+     same place a year later, on November 18 and 30, 1949. On
+     February 19, 1950, she was caught at a hollow sycamore 650
+     feet farther up the gully, and she was recaptured there on
+     February 25 and April 7, and on June 15, 1951. On August 6,
+     1951, she was caught at a house in a thicket on the gully
+     bank, between the first and second locations and 150 feet
+     from the latter.
+
+     (19.) This female was recorded only twice; on October 15,
+     1948, she was at a hilltop rock outcrop. On July 14, 1950,
+     she had moved 1480 feet and was living in a rock pile at the
+     base of the slope, near the same hollow sycamore where
+     female No. 18 had been caught.
+
+     (20.) This female was first caught as an adult on April 5,
+     1950, at a large boulder of a hillside rock outcrop. On
+     October 7, 1950, she had shifted 110 feet to a house at an
+     osage orange tree on the hilltop rock outcrop. On November 9
+     she was back at the first location and on November 28 she
+     had moved 70 feet south along the hillside outcrop. On
+     January 11 and February 9, 1951, she was back at the
+     original location. On November 9, and 21, 1951, she was
+     again at the site 70 feet south, and was still there at her
+     last capture on February 3, 1952.
+
+Ordinarily each house that is in use harbors only a single woodrat. To a
+greater degree than any other kind of mammal on this area woodrats show
+intraspecific intolerance. On various occasions when captives were
+placed in the same or adjacent cages, they focused their attention on
+each other with evident hostility, the more powerful or aggressive
+individuals attacking or pursuing. Several times the confinement of two
+rats in the same live-trap or cage resulted in the death of the weaker
+individual, and seemingly this is the normal outcome unless the attacked
+rat is able to escape. On various other occasions two or more rats have
+been caught in the same trap simultaneously but in every instance these
+were either: a pair of adults, the female appearing to be in oestrus; a
+lactating female and one or more of her young; or young less than
+half-grown, that were obviously litter mates. Older woodrats, especially
+males, often have their ears torn and punctured from fighting.
+
+Territoriality involves, primarily, defense of the house itself. An
+individual that ventures into an occupied house may be quickly routed by
+the occupant even though the latter is smaller. Chasing has been
+observed occasionally, but it is doubtful whether any individual is able
+consistently to defend the entire area over which it forages. Because
+each rat spends most of its time within the shelter of its house, an
+intruder might venture onto its home range unchallenged and undetected,
+so long as it did not enter the nest cavity.
+
+An adult female was live-trapped on October 14, 1951, beside her house
+at the outcrop. As soon as she was released, she disappeared within the
+house. After approximately two minutes, a soft, high pitched whine was
+heard and immediately another woodrat dashed into view closely followed
+by the female. The chase continued for several seconds in the vicinity
+of the house, but the woodrat being chased soon left the area _via_ the
+outcrop. Probably this intruder had moved into the house in the night
+while the female was in the trap.
+
+On June 17, 1952, an adult male was found in a live-trap set at one of
+the brush pile houses in the woodland area. This house was occupied by
+an adult female. He ran into the house after release, and immediately
+there was a loud squeal. He ran outside and paused under some limbs
+approximately 15 feet from the house, and remained there for 15 minutes
+before clipping off an ironweed 12 inches long, which he carried to the
+house. He did not enter the house but stopped beneath overhanging sticks
+at the edge, eating leaves from the plant. He made another attempt to
+enter the house but loud squeals and rustling followed and he returned
+to the ironweed plant and was still eating when observations were
+halted. In another instance, squeals and rustling indicated that the
+occupant and intruder were in combat.
+
+[Illustration: FIG. 2. Diagram illustrating spacing (due to
+territoriality or intolerance of the rats) in twelve woodrat houses in a
+hedge row extending south from south boundary of the Reservation at the
+middle.]
+
+Although home ranges may overlap to some extent, intraspecific
+intolerance tends to maintain a certain minimum interval between houses.
+The arrangement of twelve houses along a hedge row 1170 feet long is
+diagrammatically represented in Figure 2. The average interval was 78.5
+feet (minimum 42; maximum 171). The habitat was uniform. Home ranges
+probably overlap somewhat, and the spacing is the expression of the need
+for an otherwise unoccupied area in which there is sufficient space to
+live. Because individuals tend to fight whenever they meet, there is
+probably a psychological tendency for sequestration which results in
+spacing of houses and reduces social contact thereby avoiding a
+depletion of energy that would be detrimental to the population. Whereas
+condition of the hedge row determines whether or not it will be
+inhabited by woodrats, length determines the number of occupants. The
+spacing of houses in a hedge row must be attributed to something other
+than restriction of sites because the number of sites available always
+exceeds the number that are in use. Although rock outcrops situated in
+areas of uniform habitat have not been observed to the extent that hedge
+rows have, a similar spacing seems to exist and the sites available for
+houses always exceed the actual number found. This behavior pattern
+limits the number of houses and is probably advantageous to the species
+through preventing overcrowding and possible critical depletion of the
+food supply.
+
+Eleven of the young that weighed 100 grams or less when originally
+captured and were presumably still living at the mothers' houses, were
+recaptured repeatedly over periods of weeks or months, providing a
+limited amount of information regarding dispersal. They followed no
+definite pattern. In seven instances (five males and two females) the
+young stayed on at the house beyond the age when they were completely
+independent of the female. In at least two instances the female was
+known to have moved away while the young remained. One female shifted to
+a house 58 feet from the one where she had reared her litter of two, and
+was accompanied by the young male, while the young female stayed on in
+possession of the maternal house. Two months later this young female was
+caught at a house 90 feet away, and an adult male was in possession of
+her former house. One young male shifted to a house 220 feet from his
+original home and remained there several months, but was recaptured once
+back at the original location. Another male made a series of moves over
+a period of weeks and finally settled in a house 490 feet from his first
+home. One male who stayed in the maternal house all summer, and reached
+adult size there, later moved several times, and was last recorded 900
+feet away. One young female shifted 110 feet. In several instances
+juveniles appeared abruptly in houses known to have been unoccupied
+previously, and some of these houses were in poor repair. These young
+had wandered from their maternal houses, for unknown reasons. On one
+occasion a young woodrat was caught in a mouse trap set in a meadow, a
+habitat into which adult woodrats would scarcely be expected to
+venture.
+
+
+_Feeding_
+
+Rainey (1956) has listed 31 food plants that are used by the woodrat in
+northeastern Kansas. He has emphasized that each rat usually obtains its
+food from plants growing in the immediate vicinity of its house, and
+that individuals thus differ greatly in their feeding, according to the
+local vegetation. Therefore, with a sufficiently large number of
+observations, the list of food plants might be greatly expanded, to
+include most of the local flora, with the exception of the relatively
+few kinds that have developed strongly repellent properties rendering
+them unpalatable to herbivores in general.
+
+At the quarry where one or more woodrats usually lived beneath metal
+strips, as described previously (under the heading of "Commensals"), the
+situation seemed to be especially favorable, despite the fact that the
+metal offered no insulation from extremes of heat in summer and cold in
+winter. Perhaps the rat had an alternative nest among nearby boulders,
+to use when temperature was unendurable beneath the metal.
+
+The rat itself, the stored food, and other details of its home life,
+could be observed with a minimum of disturbance by raising one side of
+the metal strip momentarily, then carefully lowering it into place. The
+following observations made in the summer and autumn of 1948 give some
+idea of the range of food plants stored at any one time and the change
+as the season progresses.
+
+     July 12: Bundles of leaves of carrion-flower (_Smilax
+     herbacea_); 15 green pods of honey locust (_Gleditsia
+     triacanthos_) with seeds eaten out; several green fruits of
+     osage orange (_Maclura pomifera_), and several seeds of
+     coffee-tree (_Gymnocladus dioica_).
+
+     July 24: Bundles of green leaves of osage orange and
+     carrion-flower; many pods of honey locust.
+
+     August 30: Three large clusters of the fruits of pokeberry
+     (_Phytolacca americana_).
+
+     October 20: Many small clusters of grapes (_Vitis vulpina_)
+     judged to weigh perhaps one pound in all; several old pods
+     of coffee-tree and a few berries of dogwood (_Cornus
+     Drummondi_) and of pokeberry; a pile of small acorns of
+     chinquapin oak (_Quercus prinoides_); dry seed heads of
+     grass (_Bromus inermis_ and _B. japonicus_).
+
+     December 22: Many twigs of bittersweet (_Celastrus
+     scandens_) with fruits still attached; several seed heads of
+     sunflower (_Helianthus annuus_); a few acorns of chinquapin
+     oak; fragments of the fruit of osage orange; cured bundles
+     of trefoil (_Desmodium glutinosum_), carrion-flower, and
+     tickle grass (_Panicum capillare_).
+
+Although the eastern woodrat is relatively unspecialized in its feeding
+habits, a few species of favored food plants probably make up the
+greater part of its diet. In northeastern Kansas, at present, osage
+orange probably is by far the most important single species. Despite the
+fact that its aromatic leaves and fruits are somewhat repellent to
+insects and some other animals, they are well liked by woodrats, and
+provide a year-round food supply to those individuals having houses in
+or near the trees. Honey locust similarly provides thorny shelter for
+house sites, while the foliage, the seeds, and the bark of twigs and
+trunks are eaten. In houses that are situated near honey locusts, the
+large, heavy seed pods are sometimes stored by the hundreds. Old pods
+are often used in substitution for sticks as building material in the
+house. Nevertheless, honey locust is used relatively little as compared
+with osage orange. Other plants that figure most importantly in the diet
+include bittersweet, fox grape, pokeberry and horse nettle (_Solanum
+carolinense_).
+
+Rainey (_op. cit._) mentioned that captive woodrats would eat meat, both
+cooked and raw, and on one occasion he found remains of a cicada on a
+house under circumstances suggesting that this insect had been eaten by
+a rat. In the course of trapping for opossums and small carnivores,
+woodrats were caught on many occasions by Fitch in traps baited with
+animal material exclusively--miscellaneous meat scraps, canned dog-food,
+bacon grease, or carcasses of small vertebrates. In fact, such baits
+seemed to be even more attractive than the grain, seeds, peanut butter
+and raisins that had been used customarily to bait the traps set for
+woodrats. However, such meat baits could be used effectively only in
+cold weather, because of rapid spoilage and interference by insects at
+higher temperatures.
+
+On one occasion an adult pilot black snake found dead on the road, a
+recent traffic victim, was brought to the Reservation headquarters for
+examination and was left overnight in the garage. On the following
+morning the carcass of the snake was found to have been dragged a short
+distance and gnawed; a quantity of flesh was eaten at an exposed wound
+on the neck. Woodrat tracks were thickly imprinted on the dusty soil
+around the snake. The adult male woodrat that lived in the garage had
+evidently spent much time moving about the carcass and over it, and
+feeding upon it. It seemed remarkable that this individual was not
+deterred from feeding on the snake by an instinctive fear of one of its
+chief natural enemies.
+
+Although the eastern woodrat's food consists mostly of vegetation, the
+strong tendency noted to feed upon flesh when it is available suggests
+that these rodents may, occasionally at least, prey upon helpless young
+of small vertebrates that are readily available to them. Nestling birds,
+either on the ground or in low trees, and young mice in nests that are
+accessible, might tempt the rat to indulge in predation.
+
+
+_Breeding_
+
+Reproductive activity continues to some extent throughout the year
+except in late autumn and early winter. Presence of a vaginal orifice
+was used as an indication of sexual activity. In most instances the
+orifice was not indicative of actual oestrus, as it persisted through
+the preceding and following stages of an oestrus cycle. In anoestrus the
+orifice is sealed, the genitalia are reduced in size and the skin in the
+genital region is white. Immature females, and adults during most of the
+winter, are in this quiescent condition. Onset of the breeding season in
+late winter is relatively abrupt, and seemingly is a photoperiodic
+response. Breeding may begin in late January, and most females are in
+breeding condition within the first half of February. In oestrus the
+genitalia are enlarged and discolored and the vaginal orifice is
+prominent and gaping. By February most females born the previous season
+have matured, and breeding involves the entire population, except
+possibly for retarded young and individuals suffering from disease,
+injury or malnutrition. Rainey (1956) recorded an average of 2.3 young
+per litter.
+
+Number of litters normally produced in the course of a season by an
+adult female is unknown, but most mature females examined within the
+period February to September inclusive were in some stage of the
+breeding cycle. It is obvious that the females which are successful in
+rearing their litters produce at least two litters annually, and
+probably some produce three litters. When entire litters are lost at an
+early age, to predation, or other causes, productivity is much
+increased, with perhaps only short intervals between pregnancies.
+
+The smallest female having a vaginal orifice weighed 160 grams, but in
+most instances somewhat larger size is attained before the onset of
+oestrus. Judging from the average growth rate of immature females (Fig.
+3), most probably attain sexual maturity at an age of five to six months
+unless this age is reached in the winter period of sexual quiescence.
+Rainey (_op. cit._) found no clear cut instances of young maturing in
+time to breed before their first winter. He concluded, tentatively,
+that in most instances sexual maturity is not attained until the spring
+of the year following that in which the rat is born. However, the
+evidence was inconclusive because few of the young marked survived to
+maturity. In late summer and early autumn, the latter third of the
+breeding season, newly matured young of the year, born in early spring,
+may be the most productive group. Young conceived at the beginning of
+the breeding season, and born in early March, would normally reach adult
+size and breeding maturity in August. For example, a young female first
+caught on June 15, 1951, weighed only 150 grams, but by August 10 she
+had gained to 220 grams (probably in pregnancy) and had a vaginal
+orifice. Of 35 adult and subadult females examined by Fitch in October,
+eleven had a vaginal orifice, the latest on October 18. Of these eleven
+showing signs of breeding, four at least had not yet produced litters,
+judging from the undeveloped condition of their mammae, and others that
+showed evidence of recent lactation probably included young of the year
+that had bred in August or September. One female gave birth to a litter
+in a trap on the night of October 6, 1950. Of 32 adult and subadult
+females recorded by Fitch in November, all were sexually quiescent, with
+the possible exception of one having a partially open vagina on November
+10. All females taken in December, and most of those taken in January,
+also were sexually quiescent. January 20 was the earliest recorded date
+for a female with a vaginal orifice. Females examined in February mostly
+were perforate and many of them appeared to be in oestrus. One female
+trapped on February 19, 1950, weighed only 140 grams and was still
+imperforate. Another, weighing 200 grams on February 3, 1952, still was
+imperforate, but by February 27 she was perforate and appeared to be in
+oestrus. An adult female that appeared to be in oestrus on February 3,
+1952, was imperforate on February 10.
+
+
+_Growth_
+
+At birth woodrats weigh approximately 10 grams or a little more. In a
+litter born in captivity and kept by Rainey, the average gain amounted
+to a little more than 1.5 grams per day during the first two months, but
+in the third month it was somewhat less. As this was an unusually large
+litter, of five young, one more than the female's teats could
+accommodate, their growth may have been a little less rapid than in most
+of those under natural conditions. At an age of three months they
+averaged approximately 120 grams. The three males consistently exceeded
+the two females.
+
+[Illustration: FIG. 3. Typical growth curves for male and female
+woodrats; early stages are based on the litter of a captive female,
+later stages on average gains of recaptured juveniles and subadults,
+excluding those that seemed to be stunted. Solid line represents males
+and broken line represents females.]
+
+Young weighing less than 100 grams are rarely caught in live-traps. Four
+young, all males, first caught at an average weight of 80 grams, gained
+on the average, 1.39 grams per day over intervals that averaged 44 days.
+Six other young males first caught while in the weight range of 100 to
+149 grams, were recaptured after intervals of 17 to 45 days and they had
+gained, on the average, .92 grams per day. The corresponding figure for
+four young females in the same size range was .71 grams per day. In
+seven young males in the weight range 150 to 250 grams, that were caught
+after intervals averaging 66 days, the gain in weight amounted to .83
+grams per day. In seven females in the range 150 to 199 grams, gains
+averaged only .68 grams per day. Fully grown females that are not
+pregnant weigh, most typically, a little less than 250 grams while fully
+grown adult males average a little more than 300 grams. Growth rate and
+adult weight both are influenced to a large extent by season and even
+more by individual differences. The underlying causes are obscure in
+most instances, but individual rats that are still short of adult size
+may stop growing for periods of months, and some individuals grow much
+more rapidly than others. One male that weighed 108 grams when he was
+first caught on July 3, 1951, was estimated to have been born in early
+May. He was last captured 152 days later on December 2, 1951, and by
+then his weight was 300 grams, representing an increase of 1.2 grams per
+day. Another male that weighed only 75 grams when he was caught on
+October 8, 1950, may have been less than two months old then. By
+November 21, 1951, at a probable age of 15 months, he weighed 350 grams
+having attained almost the maximum size. Other exceptionally large
+individuals were known to be less than two years old, while those rats
+that survived longest on the study areas did not much exceed average
+adult size. These records seem to show that exceptionally large woodrats
+are usually not those of advanced age, but are individuals which have
+grown most rapidly through fortuitous circumstances, probably depending
+upon both innate and environmental factors.
+
+None of the woodrats handled was excessively fat, nor were any
+emaciated. The habit of keeping on hand stores of food at all seasons
+perhaps obviates the necessity for storing quantities of fat. Seasonal
+trends in weight vary among individuals, and are not wholly consistent
+from year to year. Rainey found that in late autumn and winter, rats
+steadily gain weight reaching a peak in late February or March. However,
+in the winters of 1948-49 and 1949-50, Fitch found that most rats lost
+weight and hardly any, even those that were short of adult size, made
+gains.
+
+The following records of a male born in the spring of 1949 show rapid
+growth and attainment of adult size in his first summer, cessation of
+growth during the winter, and resumption of growth, with attainment of
+near-maximum size the following spring.
+
+    June 16, 1949           96 gms.
+    September 26, 1949     230 gms.
+    September 27, 1949     230 gms.
+    October 18, 1949       260 gms.
+    October 27, 1949       250 gms.
+    October 29, 1949       220 gms.
+    November 8, 1949       235 gms.
+    November 15, 1949      245 gms.
+    November 24, 1949      240 gms.
+    November 26, 1949      240 gms.
+    November 30, 1949      240 gms.
+    December 20, 1949      260 gms.
+    February 18, 1950      230 gms.
+    April 5, 1950          290 gms.
+    April 7, 1950          300 gms.
+    October 7, 1950        320 gms.
+    November 29, 1950      345 gms.
+    March 23, 1951         340 gms.
+
+Another example, showing winter cessation of growth in a male at even
+smaller size is shown below. This was in the winter of 1950-1951.
+
+    November 9             145 gms.
+    November 28            175 gms.
+    November 29            165 gms.
+    January 10             180 gms.
+    January 11             175 gms.
+    March 1                225 gms.
+    March 23               200 gms.
+
+
+_Longevity_
+
+The longest span of records for an individual woodrat recorded was 991
+days in a female, already adult when she was first caught on November
+18, 1948. Other relatively long spans of records were: 827 days in a
+male, adult when first caught on March 16, 1952; 754 days in a female,
+also adult when first captured; 649 days in a male first captured as a
+juvenile; 465 days in a male, adult when first captured; 409 days in a
+male, juvenile when first captured; 399 days in a female, juvenile when
+first captured; 395 days in a female, adult when first captured; 390
+days in a female, adult when first captured; 366 days in a male, adult
+when first captured. Of these eleven individuals (six females and five
+males) whose records cover more than a year, eight were already adult
+when first caught. These eleven rats represent only 4.3 per cent of the
+total number captured. Our study was made at a time when populations
+were shrinking and disappearing, and obviously individual spans would
+have been longer if we had been working with a stable population. In
+most instances the spans of our records represent only small parts of
+the life spans of the individuals involved. Nevertheless, our records
+emphasize the potentially greater longevity of the woodrat as contrasted
+with the various smaller rodents living in the same area. Of several
+thousand individuals of the genera _Mus_, _Zapus_, _Reithrodontomys_,
+_Peromyscus_, _Sigmodon_, and especially _Microtus_, none is known to
+have survived so long as two years, and only a few individuals are known
+to have survived so long as one year after being marked.
+
+
+_Summary_
+
+Plant succession resulting from land use practices created habitat
+conditions especially favorable for woodrats in the late nineteen
+forties in northeastern Kansas, and particularly on the University of
+Kansas Natural History Reservation. With protection from prairie fires,
+woody vegetation had encroached onto areas that were formerly grassland,
+and, later, fencing against livestock permitted dense thickets of
+undergrowth to develop. In this region the woodrat usually lives in a
+forest habitat, and requires for its house sites places that are
+especially well sheltered, as in matted thickets of undergrowth, root
+tangles exposed along eroded gully banks, hollow stumps or tree trunks,
+bases of thorny trees with multiple trunks for support, thick tops of
+fallen trees, or, especially, rock outcrops with deep crevices.
+
+At the time of their maximum population density in or about 1947,
+woodrats probably averaged several per acre on the woodland parts of the
+Reservation. In the autumn of 1948, 17 were caught on the ten-acre tract
+of woodland that was live-trapped most intensively. By then, however,
+the population had already undergone drastic reduction, as shown by the
+fact that there were many unoccupied and disintegrating houses
+throughout the woodland. While the time and manner of mortality was not
+definitely determined, circumstantial evidence suggests that the
+downward trend began in early March, 1948, when record low temperatures
+and unusually heavy snowfall coincided with the time when parturition
+normally occurs. The rigorous weather conditions then may have been
+injurious, not only to the newborn litters but to the females comprising
+the breeding stock. Nevertheless, the population remained moderately
+high through 1948, but by early spring of 1949 more than three-fourths
+of the adults and subadults present in late autumn had been eliminated.
+Again, unusually severe winter weather seemed to be the underlying
+cause, as in January precipitation was the heaviest on record in 81
+years, with penetrating sleet storms, persistent ice glaze, and
+occasional brief thawing followed by sudden drops to extremely low
+temperature.
+
+After the drastic reduction in the winter of 1948-49, the population did
+not recover. Although no further sudden reductions due to extremes of
+weather were noted, the trend seemed to be one of gradual, progressive
+decline throughout the following period of years. Deterioration of the
+habitat, as the developing forest shaded out undergrowth, and inroads of
+certain predators may have been important in preventing recovery of the
+population. Many kinds of predatory mammals, hawks, owls, and snakes
+probably take woodrats occasionally, but the spotted skunk, long-tailed
+weasel, horned owl, timber rattlesnake and pilot black snake are
+considered to be by far the most important predators because of their
+habits and prey preferences. Few actual records of predation on woodrats
+were obtained because of their scarcity during most of the period
+covered by our study.
+
+Of the animals which share the woodrat's habitat, many small mammals,
+reptiles, amphibians, and invertebrates use its houses and live in a
+somewhat commensal relationship.
+
+Woodrats are somewhat territorial, each defending its house and an
+indefinite surrounding area against intrusion by others. Houses tend to
+be spaced at intervals of at least 40 feet; occasionally they are closer
+together. Most foraging for food is done within 75 feet of the house.
+However, woodrats often wander far beyond the limits of the usual home
+range. On the average, males travel more frequently and more widely than
+females, and the larger and older males travel more than the smaller and
+younger. Search for mates provides the chief motivation for wandering.
+Extent of wandering is controlled to a large degree by availability of
+natural travelways, such as rock ledges, by shelters for temporary
+stopping places, such as old deserted houses, and by population density
+of the rats themselves.
+
+Food of the eastern woodrat consists chiefly of vegetation; many kinds
+of leaves, fruits, and seeds are eaten. For many individuals foliage and
+seeds of the osage orange are the staple; hedge rows and dense trees of
+osage orange provide favorable sites for the houses. Woodrats are
+attracted to meat baits, and have been known to feed on flesh of
+carcasses, even on one of the pilot black snake which is a predator on
+the rat.
+
+Woodrats are born blind, naked, and helpless, at a weight approximately
+four per cent of the adult female's. They gain at a rate of at least 1.5
+grams per day in the first two months. When they have reached a weight
+of 100 grams, the gain averages somewhat less than one gram per day, but
+individual variation is great. Males gain more rapidly than females,
+especially in the later stages of growth, as adult weight is greater by
+approximately one-fourth in the male. Some individuals grow to maximum
+adult size at an age of one year. Unusually large individuals are not
+necessarily those that are unusually old. Longevity is greater in
+woodrats than in most smaller rodents. One female of adult size when
+first trapped was last captured 991 days later when she must have been
+well over three years old, and others are known to have survived more
+than two years even though populations were shrinking so that few of the
+rats were able to survive for their normal life span.
+
+
+
+
+_Literature Cited_
+
+
+CRABB, W. D.
+
+1941. Food habits of the prairie spotted skunk in southeastern Iowa.
+Jour. Mamm., 22:349-364.
+
+
+FITCH, H. S.
+
+1947. Predation by owls in the Sierran foothills of California. Condor,
+49:137-151.
+
+
+RAINEY, D. G.
+
+1956. Eastern woodrat, Neotoma floridana: natural history and ecology.
+Univ. Kansas Publ. Mus. Nat. Hist., 8: No. 10, in press.
+
+_Transmitted March 12, 1956._
+
+
+
+
+
+
+End of the Project Gutenberg EBook of Ecological Observations on the
+Woodrat, Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
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+
+***** This file should be named 33566-8.txt or 33566-8.zip *****
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+    <title>
+      The Project Gutenberg eBook of Ecological Observations on the Woodrat, Neotoma Floridana, by Henry S. Fitch And Dennis G. Rainey.
+    </title>
+    <style type="text/css">
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+<pre>
+
+The Project Gutenberg EBook of Ecological Observations on the Woodrat,
+Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
+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: Ecological Observations on the Woodrat, Neotoma floridana
+
+Author: Henry S. Fitch
+        Dennis G. Rainey
+
+Release Date: August 29, 2010 [EBook #33566]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK ECOLOGICAL OBSERVATIONS--WOODRAT ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci
+and the Online Distributed Proofreading Team at
+https://www.pgdp.net.
+
+
+
+
+
+
+</pre>
+
+
+
+
+<p class="center">
+<span class="smcap">University of Kansas Publications</span>
+<span class="smcap">Museum of Natural History</span><br />
+<br />
+Volume 8, No. 9, pp. 499-533, 3 figs.<br />
+<br />
+June 12, 1956<br />
+</p>
+
+<h1>Ecological Observations on the Woodrat,<br /> Neotoma floridana</h1>
+
+<h3>BY</h3>
+
+<h2>HENRY S. FITCH AND DENNIS G. RAINEY</h2>
+
+<p class="center">
+<span class="smcap">University of Kansas</span><br />
+<span class="smcap">Lawrence</span><br />
+1956<br />
+<br />
+<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br />
+<br />
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,<br />
+Robert W. Wilson<br />
+<br />
+Volume 8, No. 9, pp. 499-533<br />
+Published June 12, 1956<br />
+<br />
+<span class="smcap">University of Kansas</span><br />
+Lawrence, Kansas<br />
+<br />
+PRINTED BY<br />
+FERD VOILAND, JR., STATE PRINTER<br />
+TOPEKA, KANSAS<br />
+1956<br />
+</p>
+
+
+
+<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_501" id="Page_501">[Pg 501]</a></span></p>
+<h2>ECOLOGICAL OBSERVATIONS ON THE WOODRAT,<br /> NEOTOMA FLORIDANA</h2>
+
+<h4>By</h4>
+
+<h3>Henry S. Fitch and Dennis G. Rainey</h3>
+
+
+<h2><i>Introduction</i></h2>
+
+<p>The eastern woodrat exerts important effects on its community associates
+by its use of the vegetation for food, by providing shelter in its stick
+houses for many other small animals, and by providing a food supply for
+certain flesh-eaters. In the course of our observations on this rodent
+on the University of Kansas Natural History Reservation, extending over
+an eight-year period, from February, 1948, to February, 1956, these
+effects have changed greatly as the population of woodrats has
+constantly changed in density, and in extent of the area occupied.</p>
+
+<p>This report is concerned with the population of woodrats on the
+Reservation, the changes that the species has undergone, and the factors
+that have affected it. Our two sets of field data, used as a basis for
+this report, supplement each other and overlap little, either in time or
+space. Fitch's field work which covered approximately the western half
+of the Reservation, was begun in September, 1948, and was pursued most
+intensively in the autumn of 1948 and in 1949, with relatively small
+amounts of data obtained in 1950 and 1951 because of the great reduction
+in numbers of rats. Rainey's field work began in the spring of 1951 and
+was continued through 1954, concentrating on a colony in the extreme
+northwestern corner of the Reservation and on adjacent privately owned
+land. In actual numbers of rats live-trapped and for total number of
+records the two sets of data are comparable. Fitch's field work
+consisted chiefly of live-trapping while Rainey's relied also upon
+various other approaches to the woodrat's ecology. Rainey's findings
+were incorporated originally in a more comprehensive report (1956), from
+which short passages have been extracted that are most pertinent to the
+present discussion. Our combined data represent 258 woodrats (153
+Fitch's and 105 Rainey's) caught a total of 1110 times (660 Rainey's and
+450 Fitch's). Rainey's records pertain, in part, to woodrats outside the
+Reservation but within a few hundred yards, at most, of its boundaries.<span class='pagenum'><a name="Page_502" id="Page_502">[Pg 502]</a></span></p>
+
+
+<h2><i>Habitat</i></h2>
+
+<p>In the autumn of 1948 the population of woodrats was far below the level
+it had attained in 1947 or earlier, but the rats were still abundant and
+distributed throughout a variety of habitats. Almost every part of the
+woodland was occupied by at least a sparse population. Also, many rats
+lived beyond the limits of the woodland proper, in such places as
+deserted buildings, thickets, roadside hedges, and tangles of exposed
+tree roots along cut banks of gullies. All these situations are
+characterized by providing abundant cover, a limiting factor for this
+woodrat.</p>
+
+<p>In 1947, when the population of woodrats was especially high, plant
+succession on the wooded parts of the Reservation may have been near the
+optimum stage for the rats. For some 80 years, since the time the land
+was first settled and prairie fires were brought under control, woody
+vegetation has been encroaching into areas that were formerly grassland.</p>
+
+<p>About 1934 the University changed its policy with regard to treatment of
+the tract that was later made the Reservation. Up to that time, most of
+the area had been used as pasture and subjected to heavy grazing, but
+several fields had been fenced and cultivated. Under the new policy the
+hillsides and hilltop edges with open stands of various deciduous trees
+were enclosed with stock fences and protected from grazing. Successional
+trends were greatly altered. Woody vegetation, already favored by
+protection from the prairie fires originally important in the ecology of
+this region underwent further development as a result of protection from
+browsing. Thickets of shrubs and saplings sprang up throughout the
+woodland, forming a dense understory layer beneath the discontinuous
+canopy of the relatively scattered mature trees. The composition and
+density of the undergrowth varied markedly in different parts of the
+woodland. The parts that were formerly most open acquired the most dense
+understory. Blackberry, honey locust, osage orange, and prickly ash
+formed in places thorny tangles almost impenetrable to humans. This
+thicket stage reached its peak in density in the middle to late forties
+coinciding approximately with the time of maximum abundance of the rats.
+In the past eight years, under continued protection from burning,
+cutting and browsing, the forest has developed further; sizable trees 20
+feet or more high and up to eight inches in trunk diameter have grown
+from seedlings during the period of protection.<span class='pagenum'><a name="Page_503" id="Page_503">[Pg 503]</a></span> An almost continuous
+canopy of foliage has developed, shading the understory and thinning it
+by killing shrubs and saplings. In those situations where the canopy is
+most dense, as on north slopes having stands of young hickory averaging
+twenty feet high, the understory is now largely lacking, but in other
+situations, particularly on south slopes, the understory thickets are
+still dense. On the whole, however, habitat conditions have become less
+favorable for the woodrat.</p>
+
+<p>Within the woodland the population of woodrats was not evenly
+distributed even at its maximum density; only those situations that
+provided sufficient overhead shelter were occupied by woodrats. The
+hilltop limestone outcrop, which was the refugium of the survivors when
+the population was at low ebb, also supported the greatest concentration
+when the population was high. The number of individuals living along any
+particular stretch of ledge could be determined only by intensive
+live-trapping, whereas residences of individuals could be more readily
+identified in most other situations away from the ledge. Stick houses of
+woodrats are, characteristically, large and dome-shaped in woodland, but
+along the ledges they usually lacked this typical form and consisted of
+a much smaller accumulation of sticks, often merely filling a small
+crevice. Sticks carried into such places where they were partly or
+wholly protected from moisture and sunshine were much less subject to
+decay than those in more open situations, and remained long after the
+rats themselves were gone. Accumulations of droppings in depressions in
+rock surfaces beneath overhanging ledges likewise have lasted for many
+years. The rock outcrop provided a continuous travelway along the
+hilltops, and even parts that were not permanently occupied usually had
+some sign. The following types of situations were found to be especially
+favorable for occupancy: deep crevices beneath overhanging projections
+of the ledge; large flat boulders broken away from the main ledge; thick
+clumps of brush (usually fragrant sumac, <i>Rhus trilobata</i>) providing
+shelter and support for the house; logs fallen across the ledge
+providing support and protection for the house structure.</p>
+
+<p>A second outcropping limestone stratum approximately 20 feet below the
+level of the hilltop was just as extensive as the upper outcrop, but it
+was little used by the rats because the exposed rock surface was more
+regular, lacking the jagged cracks and deep fissures of the hilltop
+outcrop; and it lacked the overhanging projections which provided
+overhead shelter for the rats along the<span class='pagenum'><a name="Page_504" id="Page_504">[Pg 504]</a></span> upper outcrop. More than ninety
+per cent of the rats that were recorded as associated with the outcrops
+were at the hilltop stratum.</p>
+
+<p>Second in preference to the hilltop outcrop as a house site was the base
+of an osage orange tree in thick woods. This tree occurs throughout the
+woodland of the Reservation, having become established when the leaf
+canopy was more open, and the whole area was subject to grazing, with
+less development of the understory vegetation in the woodland. Houses
+were most often situated in those osage orange trees that had been cut
+one or more times, and had regenerated with spreading growth form, the
+multiple branching stems offering substantial support. Occasionally
+houses were built in crotches from two to six feet above ground.</p>
+
+<p>Blackberry thickets also are favorable locations for houses. These
+thickets grew up mostly in fenced areas from which livestock were
+excluded, but where there was not dense shade&mdash;hilltop edges and level
+or gently sloping ground adjacent to creek banks. The houses were
+usually in densest parts of the thickets where they were almost
+inaccessible. Mats of dead canes more or less horizontal, with the live
+canes growing up through them, provided effective overhead protection,
+while the ground beneath was relatively open. Houses built in the
+thickets were so well concealed that they were usually not detected
+until after leaves were shed in autumn. In most cases the blackberry
+thickets were small and well isolated. Houses of the rats were sometimes
+unusually near together suggesting that these thickets provided
+especially favorable habitat conditions.</p>
+
+<p>Hollow trees are often utilized, the accumulation of sticks for the
+house being largely inside the cavity. To be suitable for a house site,
+the snag must have an opening near ground level, and another higher on
+the trunk, providing emergency outlets in two directions. Most of the
+hollow trees utilized were black oaks (<i>Quercus velutina</i>).</p>
+
+<p>In 1948 there were many houses in cut tops of trees left from small
+scale lumbering operations a few years earlier. The densely branched
+tops of elms, oaks and hickories had satisfied the requirement for
+support of the house and nearby shelter. The houses built in them were
+in open woodland well separated from otherwise favorable situations. By
+1948 the tops were disintegrating and no longer provided effective
+shelter. The houses built in them were falling into disrepair and were
+not permanently inhabited but were often used temporarily by wandering
+individuals.<span class='pagenum'><a name="Page_505" id="Page_505">[Pg 505]</a></span></p>
+
+<p>Along cut banks of gullies where trees were partly undermined by
+erosion, the exposed, tangled root systems provided sites for occupancy.
+In these situations the accumulations of sticks were small and lacked
+the typical domed shape, consisting essentially of a lining to the
+cavity beneath the roots.</p>
+
+<p>Two small buildings at the Reservation headquarters were accessible to
+woodrats and were utilized off and on throughout much of the period of
+this study, despite the fact that most other sites of occupation away
+from the hilltop outcrops were deserted in the same period. One small
+building used as a laboratory had an enclosed wooden box five feet
+square housing an electric water pump. The interior of this box was
+accessible to the rats from beneath the floor. Litter of sticks and
+stems and various food materials were carried in by the rats. The nest
+thus protected and enclosed was not surrounded by the usual accumulation
+of sticks. An old garage 30 feet from the laboratory building was also
+occupied, sometimes by a different individual. The nest and food stores
+were behind boards propped against the wall.</p>
+
+<p>In October, 1948, live-trapping was begun on a heavily wooded slope
+facing northwest, and a ten-acre area was trapped rather thoroughly in
+the succeeding weeks. Because few traps were then available, this was
+the only area that was well sampled in 1948, although diffuse trapping
+was carried on over some 200 acres. On the ten-acre tract a total of 17
+adult and subadult woodrats were caught, four along the hilltop rock
+outcrop, six along the gully at the bottom of the slope, and seven at
+intermediate levels on the slope. Judging from the many unoccupied
+houses, the population on this tract had been much higher before the
+study was begun. On the basis of this sample it seems that in 1947 a
+population of several hundred woodrats lived on the wooded parts of the
+square mile where the Reservation is located.</p>
+
+
+<h2><i>Reduction of Population</i></h2>
+
+<p>The abrupt reduction in the population of woodrats on the Reservation
+cannot be explained conclusively with available data. Probably weather
+played a major part, but other unknown factors must have been important
+also. It is certain that the population of woodrats was high, if not at
+an all-time peak, in 1947. In late February, 1948, when one of us
+(Fitch) first visited the area on a preliminary inspection trip (not
+concerned primarily with woodrats), houses of these rats were found to
+be unusually numerous<span class='pagenum'><a name="Page_506" id="Page_506">[Pg 506]</a></span> and those seen seemed to be occupied and well
+repaired. Possibly the population was drastically reduced within the
+next few weeks, as unseasonably cold and stormy weather occurred in
+early March. For the first 12 days of March, 1948, temperature averaged
+20&deg; below that of average March weather, and even colder than the
+average for January or February. A reading of -5&deg;F. on March 11 set a
+new low locally for the month since records were begun in 1869. The
+record low temperatures were accompanied by 12.8 inches of snow. This
+spell of unusually severe weather in early March coincided with the
+period in which first litters of young usually are born, as most females
+breed in early February and the gestation period is in the neighborhood
+of five weeks. That most of these first-litter young may have been
+eliminated by the unfavorable extreme of weather at the most critical
+stage in the life cycle may be readily imagined although definite proof
+is lacking. However, the mortality must have extended beyond newborn
+young. Loss of first litters ordinarily would be compensated for by the
+end of the season, since a female usually breeds more than once in the
+course of a season. In any case, by autumn, when the actual field study
+of woodrats was initiated, many houses were already deserted and in
+disrepair. Although the rats were still moderately abundant, they were,
+seemingly, much below the population peak of the preceding year.</p>
+
+<p>Further drastic reduction of adults and subadults took place in the
+winter of 1948-49. In the course of live-trapping operations from
+mid-October into early December, 51 individuals were caught and marked.
+Chiefly because of unfavorable weather conditions, field work was
+discontinued in mid-December, and live-trapping was not resumed until
+early March. Subsequently, only 12 of the woodrats previously marked
+could be recaptured, and the population had become noticeably sparse.
+Seemingly, more than three-fourths of the population present in late
+autumn had been eliminated in the interval. In January, weather was
+exceptionally severe; on the ninth and tenth the worst sleet storm in
+twelve years occurred. Sleet fell in small granules, while the
+temperature remained several degrees below freezing. Partial thawing on
+January 12, 13 and 14 was followed by a steady drizzling rain on the
+fifteenth. On the following day the temperature dropped to -7&deg;F. Ice
+still remained from the sleet storm, and the slush again froze. On the
+night of January 18, there was one of the worst snow storms on record
+and temperature reached a low of 2&deg;F. Exceptionally low<span class='pagenum'><a name="Page_507" id="Page_507">[Pg 507]</a></span> temperatures
+persisted through January 24, with more sleet on January 25. Ice from
+the earlier storm still remained. On January 30, the temperature dropped
+to -7&deg; and a three-inch cover of snow still remained over the coat of
+ice. The month of January, 1949, had the heaviest precipitation in 81
+years (5.09 inches) and a cover of ice remained for at least 21 days.
+There were other sleet storms of lesser proportions on February 2 and
+again on February 21.</p>
+
+<p>Ordinarily sleet would not seriously damage woodrats living in houses in
+woodland habitats and less suitable hedge rows because it usually
+freezes as it falls and coats only the surface of the house. Gradual
+thawing would allow normal runoff without much penetration. Because the
+sleet during the storm described above did not form a glaze as it fell,
+the ice particles penetrated many houses. It has been observed many
+times that captive woodrats refused food that was frozen or were unable
+to eat it. Woodrats in live-traps in winter rapidly weaken unless a
+large supply of food is available. If food supplies became sealed over
+by ice, woodrats would have died by starvation or by falling an easy
+prey to predators. The rats were more accessible to several predators
+than were smaller mammals such as meadow voles which were difficult to
+obtain because of the coating of ice over the fields.</p>
+
+<p>The decimated population surviving into the breeding season of 1949
+failed to make substantial gains. In fact, during the following
+four-year period the general trend of the population over the
+Reservation as a whole seemed to be one of gradual further decline.</p>
+
+<p>In November, 1949, the rats were almost gone from the area of north
+slope and hilltop in oak-hickory-elm woodland where the most intensive
+live-trapping and other field work had been done the previous year. The
+following descriptions of houses remaining on the area at that time give
+some idea of the habitat, and of the course of events correlated with
+the fluctuations in numbers of woodrats.</p>
+
+<div class="blockquot"><p>No. 1. At the hilltop outcrop, partly on a substrate of
+limestone boulders, built around an elm of two-foot DBH,
+which lent support to one side. A hackberry sapling one inch
+in stem diameter grew through the middle of the house,
+providing further support. The house was two feet high and
+six feet in diameter, and was in obvious disrepair, with a
+hole several inches in diameter in its top. It had been
+occupied in the autumn of 1948. It was constructed mainly of
+sticks, ranging in diameter from approximately one inch to
+straw size. Many of the sticks, from .4 to .5 inches in
+diameter and one to two feet long, seemingly would have been
+heavy burdens for a rat, although they were of light-weight
+wood, sumac and elm. Mixed with the sticks were quantities
+of dry leaves, bark, and chips of wood, all material
+appearing old and weathered. This house was in
+elm-oak-hickory woods 50 feet from a cultivated field on the
+hilltop to the east and south. To the north and west the
+escarpment sloped away abruptly. There was a coralberry
+thicket beneath the trees on the adjacent hilltop.</p></div><p><span class='pagenum'><a name="Page_508" id="Page_508">[Pg 508]</a></span></p>
+
+<div class="figcenter" style="width: 600px;">
+<img src="images/fig1.jpg" width="600" height="1099" alt="" title="" />
+</div><p><span class='pagenum'><a name="Page_509" id="Page_509">[Pg 509]</a></span></p>
+
+<h4><span class="smcap">Figure 1</span></h4>
+
+<p>(A) Map of part of University of Kansas Natural History Reservation,
+showing first-capture sites for all woodrats live-trapped in the autumn
+of 1948. Because of the short time involved and the few traps available,
+much of the area shown was not thoroughly trapped. Woodrats were
+abundant, though much less so than in 1947, as shown by the large number
+of deserted houses.</p>
+
+<p>(B) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1949. Woodrats were
+still moderately abundant, but much below the level of the previous
+year. Triangles indicate those capture sites not sampled in 1948.</p>
+
+<p>(C) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1950. Numbers were
+medium-low, having undergone drastic reduction from the peak level.
+Triangles indicate those capture sites where trapping was not done in
+earlier years.</p>
+
+<p>(D) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1951. The
+population was low, but had not yet reached its lowest ebb.</p>
+
+<p>(E) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1952, when the
+population had declined to relatively low numbers and disappeared from
+much of its former habitat.</p>
+
+<p>(F) Map of the 590-acre Natural History Reservation, showing the area
+where woodrats were studied.<span class='pagenum'><a name="Page_510" id="Page_510">[Pg 510]</a></span></p>
+
+<div class="blockquot"><p>No. 2. On gently sloping hilltop edge 15 feet from the
+outcrop and escarpment, built around a forked walnut sapling
+having both trunks approximately five inches in diameter.
+The sapling, coming up through the center of the house at a
+45&deg; angle, evidently had been bent by the accumulated weight
+of the debris at an early stage of its growth, many years
+before. Trees were small in this part of the woods, with a
+well developed understory thicket of coralberry and sumac.
+This house approximately one foot high and six feet wide,
+was constructed mainly of sticks and was similar in
+composition to No. 1, but appeared considerably older with
+all the sticks blackened and rotten. In the autumn of 1948
+this house was used by woodrats, but probably only as a
+temporary stopping place, because it was already in
+disrepair then.</p>
+
+<p>No. 3. At edge of escarpment, 25 feet from No. 2, on a flat
+boulder approximately six feet long, three feet wide and one
+foot thick. The decaying and much flattened mass of sticks
+was mainly on top of the boulder, but also spilled over its
+edges. Fresh sign was noted at this house in the autumn of
+1948, but the house was already in disrepair then, and
+seemingly it was used only as a stopping place.</p>
+
+<p>No. 4. At the hilltop outcrop where an elm had fallen across
+it. The decaying log remaining was approximately 12 feet
+long and 15 inches thick. This log passed diagonally through
+the house, providing its main support. The house was
+approximately 39 inches high, its summit extending a little
+above the level of the top of the outcrop. The house was
+approximately seven feet wide along the outcrop. This house
+was somewhat intermediate between the typical dome-shaped
+stick piles that the rat builds in open situations and the
+formless accumulations of sticks with which some rats living
+in deep rock crevices line the entrances. Part of the
+accumulation was beneath the limestone boulders and
+outcropping slabs. Approximately half of the material used
+in the house consisted of sticks and the remainder of pieces
+of bark and chips of wood, mostly gathered from the fallen
+elm. This house had shrunken noticeably from decay and
+settling in the months since it was occupied, in the autumn
+of 1948. The house was surrounded by a thicket of fragrant
+sumac, dogwood, and hackberry saplings.</p>
+
+<p>No. 5. At edge of a protruding boulder one foot thick at the
+hilltop outcrop of the west facing escarpment, and 100 feet
+back in the woods from the edge of a corn field, in
+undergrowth of dogwood, wild currant, and coralberry. The
+house consisted of a pile of rotten twigs, 3 inches deep and
+30 inches wide on the upper side of the boulder, and a
+lining of similar material at the lower edge of the boulder,
+partly blocking the crevice beneath it. The twigs composing
+the house were old and rotten. However, a few dry but still
+green hackberry leaves were stored in the crevice beneath
+the boulder. In a bare space atop the boulder were several
+recent woodrat droppings, small and obviously produced by an
+immature individual, which, perhaps, had recently settled at
+this old house site.</p>
+
+<p>No. 6. In hilltop woods, 30 feet from a corner adjoining a
+pasture and a corn field, at the base of an osage orange
+tree of one foot DBH, and also over a hollow cottonwood log
+one foot in diameter and three feet from the osage<span class='pagenum'><a name="Page_511" id="Page_511">[Pg 511]</a></span> orange
+tree. Suspended mats of grape and smilax vines, and the
+thorny, dead, lower branches of the tree provided additional
+shelter. The house was composed of sticks and twigs, mostly
+of osage orange, with spines still present; slabs of bark,
+wood chips, and dry leaves also made up part of it.
+Materials on the exterior of the house appeared old and
+weathered, but the house was conical and solid. Seven fresh
+corn cobs were on the house or near its base, suggesting
+that corn from the nearby field had figured importantly in
+the diet of the occupant. A well beaten path led from the
+base of the house alongside the log, to a large cottonwood
+tree 15 feet from the house. This evidence that the house
+was occupied was verified by live-trapping the occupant.
+Late in 1948, also, the house was occupied by another
+individual, but seemingly was deserted for a period of
+months thereafter.</p>
+
+<p>No. 7. On upper part of north slope where a hickory seven
+inches in diameter had fallen across an old sunken log
+approximately one foot in diameter. The house, composed
+mainly of hickory twigs 1/4-inch to 1/2-inch in diameter,
+mixed with bark, wood chips, and leaves, was partly decayed,
+with no fresh sign and was in a thicket of greenbrier,
+saplings of hickory and hackberry, and cut tops of
+hickories. The top was flattened to less than four inches
+above the level of the supporting hickory log. There were
+large cavities in the side of the house. When first
+discovered in the autumn of 1948, this house was occupied by
+a subadult female rat, but she moved away permanently, and
+the house had been deserted for approximately a year when
+these observations were recorded.</p>
+
+<p>No. 8. In middle of northwest slope, in thick branches of
+broken top of a black oak. This house had become flattened
+by decay and settling to form a mound approximately one foot
+high and five feet in diameter. Only the top protruded
+through the carpet of dry leaves. Once well protected and
+partly concealed by the branches and twigs of the oak top,
+this house was now fully exposed by the disintegration of
+the top. The house consisted chiefly of oak twigs. In
+October, 1948, a woodrat was live-trapped at this house, but
+probably it was a wanderer. The house had then already
+undergone much deterioration.</p></div>
+
+
+<h2><i>Natural Enemies</i></h2>
+
+<p>Some 56 species of animals that regularly prey on small vertebrates live
+on the Reservation. Many of the larger kinds may take woodrats
+occasionally. Because of size, habitat preferences and the time and
+manner of hunting, five species stand out as the more formidable
+enemies&mdash;the horned owl (<i>Bubo virginianus</i>), prairie spotted skunk
+(<i>Spilogale putorius</i>), long-tailed weasel (<i>Mustela frenata</i>), pilot
+black snake (<i>Elaphe obsoleta</i>) and timber rattlesnake (<i>Crotalus
+horridus</i>).</p>
+
+<p>Throughout the study horned owls were common on the area, but their
+numbers were highest in 1948. Samples of pellet collections have shown
+that the cottontail is the staple food, being represented in almost
+every pellet. Various rodents also are important in the diet, the cotton
+rat, prairie vole, or white-footed mouse being<span class='pagenum'><a name="Page_512" id="Page_512">[Pg 512]</a></span> most prominent according
+to the time and place of collection. The woodrat is approximately
+optimum size for prey, and it constitutes one of the most preferred food
+sources. Remains of only two woodrats were found in the pellets
+examined, but at times when the pellets were collected woodrats were so
+scarce that they constituted only an insignificant percentage of the
+biomass of potential prey. On several occasions woodrats in live-traps
+were attacked by horned owls, as shown by the overturned and displaced
+trap and quantities of fine down adhering to them and to nearby objects.
+The horned owl lives in the same habitat as does the woodrat. In other
+regions woodrats are known to figure prominently in the diet of the
+horned owl. At the San Joaquin Experimental Range in California, for
+instance, <i>N. fuscipes</i> was found 240 times, more frequently than any
+other kind of prey, in 654 pellets of the horned owl, and this owl was
+shown to be the one most important natural enemy of the rat, although
+many kinds of carnivores, raptors and snakes also took toll from its
+populations. On the Reservation the population of horned owls has been
+fairly stable from year to year, with roughly one pair to 100 acres of
+woodland. Some territories have been maintained continuously throughout
+the eight-year period of observation, though changing to some extent in
+size, shape and area included. In 1948, when livestock grazed on the
+area, and the ground cover of herbaceous vegetation was relatively
+sparse, cottontails were much less abundant than they were later when
+the vegetation was protected. Small rodents including voles, cottonrats,
+and deer mice, were also less abundant then, and the numerous horned
+owls may have been supported in part by the high population of woodrats.</p>
+
+<p>The spotted skunk may be an even more important enemy of the woodrat,
+although the evidence is circumstantial. No records of these skunks
+preying on woodrats have been found in the literature, nor were any such
+instances recorded by us except for attacks on woodrats confined in
+live-traps. This skunk is a formidable enemy of small and medium-sized
+rodents, as it can climb, dig, and squeeze through small openings. That
+it may prey on rat-sized rodents and may even be a limiting factor to
+their occurrence is well shown by Crabb's (1941:353) studies in Iowa. He
+found that Norway rats (<i>Rattus norvegicus</i>) ranked third in frequency
+(cottontail, mostly carrion, ranked first) in the winter food of the
+spotted skunk. Crabb observed that about farmyards and farm buildings
+where the skunks had been eliminated by persistent<span class='pagenum'><a name="Page_513" id="Page_513">[Pg 513]</a></span> persecution, rats
+were abundant, but that about others where the skunks were present, the
+rats were scarce or absent. On several occasions he noted that heavy
+populations of rats about farm buildings in summer and autumn nearly
+disappeared in winter if a skunk was in residence.</p>
+
+<p>Sign of spotted skunk was noted frequently on various parts of the
+Reservation, especially along the hilltop ledges which were the best
+woodrat habitat. On several occasions skunks released from live-traps
+took shelter in woodrat houses which appeared to be unoccupied.
+According to a local fur dealer, C. W. Ogle, spotted skunks reached a
+peak of abundance in Douglas County in the winter of 1947-1948, and many
+pelts were brought in for sale then. The concentration of skunks may
+have had detrimental effect on the population of woodrats, especially
+when extremes of weather had already made conditions critical for them,
+as in early March, 1948, and in January, 1949, when snow and sleet made
+their usual food supply unavailable.</p>
+
+<p>The long-tailed weasel is considered to be a potentially important enemy
+of the woodrat. Weasels have been seen on the Reservation on only a few
+occasions, but they may be more numerous than these records would
+indicate. Two were caught at the hilltop outcrop, at different times and
+places, in funnel traps put out to catch snakes. The weasel seems to
+prefer this rocky habitat, which is also favored by the woodrat. Because
+of its ferocity and willingness to attack relatively large prey, and
+because it is an agile climber and able to squeeze through any openings
+large enough to accommodate a woodrat, it would seem to be a formidable
+enemy.</p>
+
+<p>The pilot black snake (<i>Elaphe obsoleta</i>) is an important enemy of this
+woodrat on the Reservation and probably throughout the rat's geographic
+range except for the extreme western part. Although this snake occurs in
+every habitat of the Reservation, it has been found most often along
+rock outcrops of wooded hilltop edges in the type of habitat most
+favored by the rat. Most often pilot black snakes have attempted to
+escape into crevices of the outcrop. These snakes are also skillful
+climbers and often have escaped by climbing out of reach along branches
+or even vertical tree trunks. On several occasions these snakes have
+been found on or beside woodrat houses, or have escaped into them. Over
+a seven-year period 143 pilot black snakes have been recorded, 53 of
+which were adults.<span class='pagenum'><a name="Page_514" id="Page_514">[Pg 514]</a></span></p>
+
+<p>On September, 1948, a large pilot black snake found basking on a rock
+ledge, distended by a recent meal, was palped and contained a subadult
+female woodrat. On June 19, 1953, one of us, approaching a live-trap set
+under an overhanging rock ledge, saw a four-foot pilot black snake on
+top of it. The snake struck repeatedly at the rat in this trap, but was
+unable to reach it. At each stroke the rat would dash about the trap
+frantically.</p>
+
+<p>These snakes hunt by stealth, and might catch woodrats by entering their
+nests, or by lying in wait along their runways, but are not quick enough
+to catch them in actual pursuit. Young in the nest would seem to be
+especially susceptible to predation by the pilot black snake. These
+snakes hunt by active prowling, either by night or by day, and much of
+their food consists of the helpless young of birds and mammals found in
+the nests. While only well-grown or adult pilot black snakes would be
+able to swallow an adult woodrat, any but first-year young probably
+would be able to overcome and swallow the small young. The female
+woodrat's habit of dragging the young attached to her teats as she flees
+from the house at any alarm must save many litters from predation by the
+pilot black snake. First litters of young, born in early March, are
+already well grown, and past the age of greatest susceptibility to
+predation before the snakes emerge from hibernation in late April or
+early May.</p>
+
+<p>The timber rattlesnake is another potentially destructive enemy, but on
+the Reservation, and throughout much of its original range it is now
+relatively scarce. The genus <i>Neotoma</i> largely coincides in its over-all
+distribution with the genus <i>Crotalus</i>, of the rattlesnakes. For most
+kinds of woodrats, the larger species of rattlesnakes are among the
+chief natural enemies.</p>
+
+<p>The timber rattlesnake has habitat preferences similar to those of the
+eastern woodrat. Of 30 timber rattlesnakes recorded on the Reservation
+over an eight-year period, all but one were at or near hilltop rock
+ledges in woodland. The woodrat is probably one of the most important
+prey species for the timber rattlesnake. Like the woodrat, the
+rattlesnake is mostly nocturnal in its activity. Unlike the pilot black
+snake, it hunts by lying in wait, striking prey which comes within
+range, and waiting for it to die from the venomous bite, rather than by
+active prowling. Therefore, it is probably less of a hazard to young in
+the nest than is the pilot black snake. Even young rattlesnakes too
+small to eat woodrats are potentially dangerous to them, as they may
+strike and kill any that come within range.<span class='pagenum'><a name="Page_515" id="Page_515">[Pg 515]</a></span></p>
+
+
+<h2><i>Commensals</i></h2>
+
+<p>Rainey (1956) listed many kinds of small animals that use the houses of
+the eastern woodrat and live in more or less commensal relationships
+with these rodents.</p>
+
+<p>A situation unusually favorable for observing woodrats and their
+associates was discovered on the Reservation where, in July, 1948, two
+old strips of sheet metal, each covering an area of approximately 25
+square feet, were used as shelter by a lactating female with three
+young. This was on a brushy slope just below an old quarry site. A rock
+pile and remains of an old rock wall were nearby. Woodrats had carried
+many sticks back under the metal strips, filling the spaces beneath
+their edges. There was a nest and a system of runways beneath the
+strips. In the following seven years this site was seldom deserted for
+long and was used by a succession of individuals. The strips of metal
+could be easily raised and then lowered into place with little
+disturbance. Because the situation was not entirely natural, the
+findings may not be typical of other rat houses. Animals found over a
+period of years beneath these metal strips include: several dozen each
+of the ring-necked snake (<i>Diadophis punctatus</i>), five-lined skink
+(<i>Eumeces fasciatus</i>), and ant-eating toad (<i>Gastrophryne olivacea</i>);
+several individuals each of cottontail (<i>Sylvilagus floridanus</i>),
+white-footed mouse (<i>Peromyscus leucopus</i>), short-tailed shrew (<i>Blarina
+brevicauda</i>), least shrew (<i>Cryptotis parva</i>), American toad (<i>Bufo
+americanus</i>), Great Plains skink (<i>Eumeces obsoletus</i>), pilot black
+snake (<i>Elaphe obsoleta</i>); and one each of bull snake (<i>Pituophis
+catenifer</i>), spotted king snake (<i>Lampropeltis calligaster</i>), red milk
+snake (<i>L. triangulum</i>), and timber rattlesnake (<i>Crotalus horridus</i>).
+The snakes which were potential predators on the rats seemed to be
+merely utilizing the shelter in these instances, but they may have been
+lying in wait for prey there.</p>
+
+<p>Among mammals, the cottontail and the white-footed mouse are the most
+persistent users of the woodrat houses, especially those that are no
+longer occupied by the rats. On one occasion five white-footed mice were
+caught simultaneously in a trap set beside a house at the base of an
+osage orange tree. Subsequent trapping showed that this house was no
+longer occupied by a rat, but that the mice lived in it. Occupancy of
+such an old woodrat house by white-footed mice may continue long after
+abandonment of the house by the rat, even after the house has partly
+decayed and settled to a small part of its original volume.<span class='pagenum'><a name="Page_516" id="Page_516">[Pg 516]</a></span></p>
+
+<p>Cottontails often have their forms under the edges of houses, either
+occupied or deserted. These situations offer protection overhead and on
+three sides. Abandoned houses having one or more of the entrance holes
+enlarged, as by predators breaking through the side of the house to gain
+access to the nest, are especially well adapted for occupancy by the
+cottontail. The rabbit may make its form inside the house structure.</p>
+
+<p>The opossum, also, finds the type of shelter that it requires in
+abandoned houses that have had the entrances sufficiently enlarged. On
+various occasions opossums or their remains have been found in such old
+houses, and opossums released from live-traps have been known to seek
+shelter in abandoned woodrat houses.</p>
+
+<p>At the old quarry on the Reservation woodrat sign was especially
+abundant. A wooden bin approximately seven feet square, used to store
+crushed rock before quarrying operations were abandoned, was inhabited
+by one rat. At the base of a rock crusher on the top of a bank a few
+yards from the bin was an accumulation of sticks and other debris
+brought by woodrats. A rock wall at the top of the bank between the
+crusher and the bin had many crevices providing shelter for the rats,
+and projecting rocks were littered with their droppings. In the spring
+of 1949 the bin and rock crusher were removed, but at least one rat
+continued to live in the rock wall. In the summer of 1951 several tons
+of corn ruined in the flood were dumped on the top of the bank above the
+wall. By autumn, Norway rats, either brought in with the corn or
+attracted by it, had taken possession of the wall, evidently displacing
+the woodrats, which were no longer present. Although this Old World
+murid rat is much different from the woodrat in habits, it seemingly can
+compete with it and replace it where habitat conditions are otherwise
+favorable for both.</p>
+
+
+<h2><i>Movements</i></h2>
+
+<p>The woodrat is dependent on the stick houses that it constructs for
+shelter. For each individual the house constitutes a home base to which
+it is attached, and about which its movements revolve. The area within
+which routine daily movements are confined constitutes the home range,
+which is variable in size and shape. An individual may, and usually
+does, alter its home range over periods of time. The home range is
+somewhat nebulous because the rat may at any time move far beyond the
+small area to which its activities are largely confined. It may be
+motivated by sexual urge or other<span class='pagenum'><a name="Page_517" id="Page_517">[Pg 517]</a></span> voluntary wandering; it may be
+enticed by a food supply or some other specific attraction not available
+near its house; or it may be forcibly displaced by an intruder or may
+abandon in favor of an offspring.</p>
+
+<p>An occupied house normally has several runways radiating from it. These
+are well worn paths, smoothed by use, and cleared of obstructions, and
+the rat tends to keep to them in its foraging expeditions. Usually a
+trail leads to a bush or tree showing evidence of heavy use by the rat.
+Ordinarily such a trail cannot be traced more than 30 feet from the
+house, and it seems that the most concentrated foraging occurs within
+this short radius. Experience in live-trapping has indicated that the
+distance covered by a woodrat in its normal foraging for food is
+ordinarily less than 75 feet in any direction from the house.</p>
+
+<p>Usually the rats can be caught in traps only at their houses or nearby
+places that they frequent, as indicated by their sign. When travelling,
+woodrats make use of overhead cover as much as possible. Storing of food
+seems to be associated with the animal's reluctance to wander far from
+home. When a rat is gathering preferred food for storage the home range
+may be enlarged (or the animal may travel beyond the limits of its
+regular home range). In any case the rat may find it necessary to
+traverse an additional area in order to reach the food source. This may
+involve, in part, extension vertically, as when the rat obtains food
+from trees directly over the house. The home range is thus somewhat
+three-dimensional; both trails and feeding places are often above
+ground. Because of dependency on cover, woodrats do not forage randomly
+in all directions from the house.</p>
+
+<p>Although the house and its immediate environs are defended as a
+territory by the occupant, possession may be soon relinquished. A
+woodrat may shift frequently from one house to another, especially if
+unoccupied houses are readily available. Because woodrats had undergone
+drastic reduction in numbers, as discussed on p. 505, unoccupied houses
+in various stages of disrepair were numerous throughout the woodland in
+1948 and 1949, and the rats that were present then seemed especially
+inclined to wander. Even old houses that are collapsed and
+disintegrating may be used temporarily, or may be taken over and
+repaired. Houses that are in sites exceptionally favorable in that they
+provide food and shelter may be occupied more or less permanently, with
+a succession of woodrats over many generations.<span class='pagenum'><a name="Page_518" id="Page_518">[Pg 518]</a></span></p>
+
+<p>Shifts to new areas are perhaps most often motivated by a search for
+mates. Such shifts are, on the average, longer and more frequent in
+males. Males must range farther in search of females when numbers are
+low. On the other hand, when numbers are high and most of the best sites
+are occupied, newly independent young and displaced adults are forced to
+travel greater distances in search of homes. Some of the larger and more
+powerful males move far greater distances than smaller males. The
+longest distances recorded were mostly for large adult males in breeding
+condition. The average maximum distance between successive points of
+capture for 27 adult males was 345 feet. For 39 females (adults and
+subadults) the corresponding figure was 143 feet. The extremes for males
+were 0 to 1080 feet and for females, 0 to 650 feet. Of the 27 males,
+five moved the maximum distance in a single night. Most of the long
+movements by males did not constitute clear-cut shifts in home range,
+and many returned to their original locations.</p>
+
+<p>The average distance between points of first and last captures for 72
+subadult and adult males was 165 feet. A similar figure for 72 subadult
+and adult females was 133 feet. Of the males 23.7 per cent were at the
+same place at the first and last captures; for females the percentage
+was 36.1. These figures are from the combined data of our trapping
+records, but the trends differed sharply in the two sets of records. In
+Fitch's records, movements averaged longer and difference between the
+sexes was much less: 189 feet for 41 males and 178 feet for 42 females.
+Corresponding figures from Rainey's records were: 141 feet for 31 males
+and 74 feet for 30 females. In Fitch's field work, opportunities to
+record exceptionally long movements obviously were better because the
+trap line encompassed a larger area, approximately half a square mile,
+whereas Rainey's live-trapping was concentrated on relatively small
+areas. The reason for the greater vagility of females in Fitch's records
+is less evident. However, the data were obtained within the period of
+drastic population reduction, at a time when there were numerous empty
+houses throughout the woodland, facilitating travel, and shifts from one
+home range to another where conditions were, temporarily at least, more
+favorable. Rainey found that the females in the small colony in woodland
+where he trapped, moved much less than did those that lived along the
+hilltop outcrop, which provided a natural travel route.</p>
+
+<p>Following are several examples of males and females with long histories
+showing individual variation in frequency and distance of movements.<span class='pagenum'><a name="Page_519" id="Page_519">[Pg 519]</a></span></p>
+
+<div class="blockquot"><p class="center"><i>Males</i></p>
+
+<p>(1.) First captured October 14, 1951, and last captured 327
+days later on September 6, 1952. He was taken 12 times. For
+the first seven captures (October 14, 1951, to July 15,
+1952), no movements were recorded. In the following seven
+days he moved 367 feet. Within the next 21 days he returned
+to within 114 feet of the site of original capture. Less
+than one month later he was caught for the last time, at
+this same site.</p>
+
+<p>(2.) This large male was captured twelve times over a period
+of 827 days (March 16, 1952, to June 21, 1954). He tended to
+wander more than other males and was absent from the
+trapping area from early 1952 to May 1953. One round trip
+made in a two-weeks period, amounted to a linear distance of
+1894 feet if the rat followed natural cover. The return trip
+of 947 feet was the greatest distance traversed in a single
+night in any of the woodrats we recorded. Other movements
+between successive captures were: 722, 397, 356, 293, 253
+and 144 feet (the latter shift made three different times).
+Sexual urge probably motivated most of his wandering, since
+numbers of females were low.</p>
+
+<p>(3.) For this male the span of records was 143 days, with 18
+captures. For the first eight recaptures, extending over a
+period of 39 days, he was still at the original location.
+Four days later he had moved 120 feet and was visiting a
+female. A week later he returned. In the following month he
+was recorded as making two more moves, of 115 feet and 215
+feet. He was last recorded at the hilltop outcrop.</p>
+
+<p>(4.) The records of this male extended over 465 days, with
+13 captures. For the entire period only one movement, of 163
+feet, was recorded. Twelve of the 13 captures were at the
+same house.</p>
+
+<p>(5.) This male was captured 16 times over a span of 130
+days. After the second capture he moved 144 feet along the
+outcrop and was caught there for the next 14 times, having
+developed a "trap habit."</p>
+
+<p>(6.) This male was in the area 210 days (13 captures) and
+shifted his range. He was first captured on August 17, 1952,
+at a house at the rock fence 433 feet from the outcrop.
+Between this date and October 12, 1952, he moved to the
+outcrop and established residence in a vacant house. He was
+recorded as making six more moves, the longest of which was
+only 40 feet.</p>
+
+<p>(7.) This male was first caught in June, 1949, as a juvenile
+probably between two and three months old (weighing 96
+grams) and hence probably still at the maternal house. In
+September, grown to adult size, he was caught twice, still
+at this same place. In October, November, December, and in
+February, 1950, he was caught 11 times at eight places all
+within a 90-foot radius of his original location. In April,
+1950, he was caught at points 550 feet WSW and 700 feet SW.
+In October he was caught within 150 feet of the original
+location. In November, 1950, and in March and April, 1951,
+he was caught four times at a place 900 feet SW from his
+original location.</p>
+
+<p>(8.) This subadult male was first caught at the hilltop
+outcrop on October 4, 1949. Two days later he had moved 160
+feet north along the outcrop. A month later he had shifted
+600 feet south; in three more days 1040 feet north. On
+November 15 he was 105 feet south of the November 8
+location; on November 16, he had moved 70 feet north. On
+November 17 he had moved 900 feet back south, but had
+returned on the 18th to the November 16 location. On<span class='pagenum'><a name="Page_520" id="Page_520">[Pg 520]</a></span>
+November 22, he had again shifted 900 feet south. All
+capture sites were at the hilltop outcrop.</p>
+
+<p>(9.) This male was caught as a juvenile (75 grams) on
+October 8, 1950. On November 9 he had moved 220 feet, from
+the lower outcrop to the upper, and he was recaptured at or
+near this same site on November 10, 28 and 29, and on
+January 11 and February 9, 1951. On November 21, 1951, grown
+to maximum adult size, he was caught at a new location 1080
+feet from the original.</p>
+
+<p>(10.) This male was caught as a subadult twice at the same
+place on November 30 and December 14. By the following
+autumn he had shifted to a new location 180 feet south along
+the outcrop, and he was caught there on September 22 and
+October 18, 1951, and on January 20 and February 2, 1952.</p>
+
+
+<p class="center"><i>Females</i></p>
+
+<p>(11.) This female was captured 27 times over a span of 211
+days. She moved back and forth considerably between two
+houses 40 feet apart but made only one substantial movement
+of 245 feet; at this time she was in breeding condition.
+Nearly seven months after the first capture she was seen for
+the last time only 16 feet from the original site of
+capture. It was assumed she fell prey to spotted skunks
+which were raiding traps.</p>
+
+<p>(12.) First captured on March 24, 1951, she remained on the
+area 105 days in which period she was live-trapped 25 times.
+Sixty per cent of the total captures were at the same house
+and the longest movement recorded was only 56 feet. She was
+last caught in a trap 25 feet from the site of original
+capture.</p>
+
+<p>(13.) This young adult remained at her house at the rock
+fence approximately four months. In this period she was
+captured 11 times. On March 16, 1952, she had moved 410 feet
+to a house at the eastern section of outcrop, probably
+searching for a male. She was never seen again.</p>
+
+<p>(14.) This subadult female moved from the site of original
+capture to a house 253 feet away on the same outcrop. She
+was probably in search of a new home when caught the first
+time. She was recorded at another house 40 feet away on one
+occasion.</p>
+
+<p>(15.) Over a span of 90 days and 15 captures this female was
+not recorded as making any movement. She was living in one
+of the woodland houses. Mature males were numerous in the
+area and she was visited by at least two.</p>
+
+<p>(16.) This female was also living in the woodland section
+and was first caught on March 30, 1952, in one of the less
+favorable houses. She was trapped 17 times over a period of
+85 days. One movement of 68 feet to a new home site was
+recorded, but the area of foraging probably did not change.
+She was caught here four times and then disappeared.</p>
+
+<p>(17.) This female was first trapped as a subadult on October
+5, 1948, at a house in brush on the upper part of a north
+slope. On November 24 she had shifted 590 feet to the bottom
+of the slope and was living in the recess beneath an
+undermined honey locust on a gully bank. On November 25 she
+was caught in a similar situation 100 feet farther east
+along the gully bank. She was recaptured at the gully on
+November 26 and 30, December 1, 3, 22, and March 8 and 9,
+and in all she shifted six times between the two gully-bank
+dens.<span class='pagenum'><a name="Page_521" id="Page_521">[Pg 521]</a></span></p>
+
+<p>(18.) This female was first trapped as an adult on November
+18, 1948, in a gully-bank den. She was recaptured at this
+same place a year later, on November 18 and 30, 1949. On
+February 19, 1950, she was caught at a hollow sycamore 650
+feet farther up the gully, and she was recaptured there on
+February 25 and April 7, and on June 15, 1951. On August 6,
+1951, she was caught at a house in a thicket on the gully
+bank, between the first and second locations and 150 feet
+from the latter.</p>
+
+<p>(19.) This female was recorded only twice; on October 15,
+1948, she was at a hilltop rock outcrop. On July 14, 1950,
+she had moved 1480 feet and was living in a rock pile at the
+base of the slope, near the same hollow sycamore where
+female No. 18 had been caught.</p>
+
+<p>(20.) This female was first caught as an adult on April 5,
+1950, at a large boulder of a hillside rock outcrop. On
+October 7, 1950, she had shifted 110 feet to a house at an
+osage orange tree on the hilltop rock outcrop. On November 9
+she was back at the first location and on November 28 she
+had moved 70 feet south along the hillside outcrop. On
+January 11 and February 9, 1951, she was back at the
+original location. On November 9, and 21, 1951, she was
+again at the site 70 feet south, and was still there at her
+last capture on February 3, 1952.</p></div>
+
+<p>Ordinarily each house that is in use harbors only a single woodrat. To a
+greater degree than any other kind of mammal on this area woodrats show
+intraspecific intolerance. On various occasions when captives were
+placed in the same or adjacent cages, they focused their attention on
+each other with evident hostility, the more powerful or aggressive
+individuals attacking or pursuing. Several times the confinement of two
+rats in the same live-trap or cage resulted in the death of the weaker
+individual, and seemingly this is the normal outcome unless the attacked
+rat is able to escape. On various other occasions two or more rats have
+been caught in the same trap simultaneously but in every instance these
+were either: a pair of adults, the female appearing to be in oestrus; a
+lactating female and one or more of her young; or young less than
+half-grown, that were obviously litter mates. Older woodrats, especially
+males, often have their ears torn and punctured from fighting.</p>
+
+<p>Territoriality involves, primarily, defense of the house itself. An
+individual that ventures into an occupied house may be quickly routed by
+the occupant even though the latter is smaller. Chasing has been
+observed occasionally, but it is doubtful whether any individual is able
+consistently to defend the entire area over which it forages. Because
+each rat spends most of its time within the shelter of its house, an
+intruder might venture onto its home range unchallenged and undetected,
+so long as it did not enter the nest cavity.<span class='pagenum'><a name="Page_522" id="Page_522">[Pg 522]</a></span></p>
+
+<p>An adult female was live-trapped on October 14, 1951, beside her house
+at the outcrop. As soon as she was released, she disappeared within the
+house. After approximately two minutes, a soft, high pitched whine was
+heard and immediately another woodrat dashed into view closely followed
+by the female. The chase continued for several seconds in the vicinity
+of the house, but the woodrat being chased soon left the area <i>via</i> the
+outcrop. Probably this intruder had moved into the house in the night
+while the female was in the trap.</p>
+
+<p>On June 17, 1952, an adult male was found in a live-trap set at one of
+the brush pile houses in the woodland area. This house was occupied by
+an adult female. He ran into the house after release, and immediately
+there was a loud squeal. He ran outside and paused under some limbs
+approximately 15 feet from the house, and remained there for 15 minutes
+before clipping off an ironweed 12 inches long, which he carried to the
+house. He did not enter the house but stopped beneath overhanging sticks
+at the edge, eating leaves from the plant. He made another attempt to
+enter the house but loud squeals and rustling followed and he returned
+to the ironweed plant and was still eating when observations were
+halted. In another instance, squeals and rustling indicated that the
+occupant and intruder were in combat.</p>
+
+<div class="figcenter" style="width: 650px;">
+<img src="images/fig2.jpg" width="650" height="151" alt="Fig. 2. Diagram illustrating spacing (due to
+territoriality or intolerance of the rats) in twelve woodrat houses in a
+hedge row extending south from south boundary of the Reservation at the
+middle." title="" />
+<span class="caption">Fig. 2. Diagram illustrating spacing (due to
+territoriality or intolerance of the rats) in twelve woodrat houses in a
+hedge row extending south from south boundary of the Reservation at the
+middle.</span>
+</div>
+
+<p>Although home ranges may overlap to some extent, intraspecific
+intolerance tends to maintain a certain minimum interval between houses.
+The arrangement of twelve houses along a hedge row 1170 feet long is
+diagrammatically represented in Figure 2. The average interval was 78.5
+feet (minimum 42; maximum 171). The habitat was uniform. Home ranges
+probably overlap somewhat, and the spacing is the expression of the need
+for an otherwise unoccupied area in which there is sufficient space to
+live. Because individuals tend to fight whenever they meet, there is
+probably a psychological<span class='pagenum'><a name="Page_523" id="Page_523">[Pg 523]</a></span> tendency for sequestration which results in
+spacing of houses and reduces social contact thereby avoiding a
+depletion of energy that would be detrimental to the population. Whereas
+condition of the hedge row determines whether or not it will be
+inhabited by woodrats, length determines the number of occupants. The
+spacing of houses in a hedge row must be attributed to something other
+than restriction of sites because the number of sites available always
+exceeds the number that are in use. Although rock outcrops situated in
+areas of uniform habitat have not been observed to the extent that hedge
+rows have, a similar spacing seems to exist and the sites available for
+houses always exceed the actual number found. This behavior pattern
+limits the number of houses and is probably advantageous to the species
+through preventing overcrowding and possible critical depletion of the
+food supply.</p>
+
+<p>Eleven of the young that weighed 100 grams or less when originally
+captured and were presumably still living at the mothers' houses, were
+recaptured repeatedly over periods of weeks or months, providing a
+limited amount of information regarding dispersal. They followed no
+definite pattern. In seven instances (five males and two females) the
+young stayed on at the house beyond the age when they were completely
+independent of the female. In at least two instances the female was
+known to have moved away while the young remained. One female shifted to
+a house 58 feet from the one where she had reared her litter of two, and
+was accompanied by the young male, while the young female stayed on in
+possession of the maternal house. Two months later this young female was
+caught at a house 90 feet away, and an adult male was in possession of
+her former house. One young male shifted to a house 220 feet from his
+original home and remained there several months, but was recaptured once
+back at the original location. Another male made a series of moves over
+a period of weeks and finally settled in a house 490 feet from his first
+home. One male who stayed in the maternal house all summer, and reached
+adult size there, later moved several times, and was last recorded 900
+feet away. One young female shifted 110 feet. In several instances
+juveniles appeared abruptly in houses known to have been unoccupied
+previously, and some of these houses were in poor repair. These young
+had wandered from their maternal houses, for unknown reasons. On one
+occasion a young woodrat was caught in a mouse trap set in a meadow, a
+habitat into which adult woodrats would scarcely be expected to
+venture.<span class='pagenum'><a name="Page_524" id="Page_524">[Pg 524]</a></span></p>
+
+
+<h2><i>Feeding</i></h2>
+
+<p>Rainey (1956) has listed 31 food plants that are used by the woodrat in
+northeastern Kansas. He has emphasized that each rat usually obtains its
+food from plants growing in the immediate vicinity of its house, and
+that individuals thus differ greatly in their feeding, according to the
+local vegetation. Therefore, with a sufficiently large number of
+observations, the list of food plants might be greatly expanded, to
+include most of the local flora, with the exception of the relatively
+few kinds that have developed strongly repellent properties rendering
+them unpalatable to herbivores in general.</p>
+
+<p>At the quarry where one or more woodrats usually lived beneath metal
+strips, as described previously (under the heading of "Commensals"), the
+situation seemed to be especially favorable, despite the fact that the
+metal offered no insulation from extremes of heat in summer and cold in
+winter. Perhaps the rat had an alternative nest among nearby boulders,
+to use when temperature was unendurable beneath the metal.</p>
+
+<p>The rat itself, the stored food, and other details of its home life,
+could be observed with a minimum of disturbance by raising one side of
+the metal strip momentarily, then carefully lowering it into place. The
+following observations made in the summer and autumn of 1948 give some
+idea of the range of food plants stored at any one time and the change
+as the season progresses.</p>
+
+<div class="blockquot"><p>July 12: Bundles of leaves of carrion-flower (<i>Smilax
+herbacea</i>); 15 green pods of honey locust (<i>Gleditsia
+triacanthos</i>) with seeds eaten out; several green fruits of
+osage orange (<i>Maclura pomifera</i>), and several seeds of
+coffee-tree (<i>Gymnocladus dioica</i>).</p>
+
+<p>July 24: Bundles of green leaves of osage orange and
+carrion-flower; many pods of honey locust.</p>
+
+<p>August 30: Three large clusters of the fruits of pokeberry
+(<i>Phytolacca americana</i>).</p>
+
+<p>October 20: Many small clusters of grapes (<i>Vitis vulpina</i>)
+judged to weigh perhaps one pound in all; several old pods
+of coffee-tree and a few berries of dogwood (<i>Cornus
+Drummondi</i>) and of pokeberry; a pile of small acorns of
+chinquapin oak (<i>Quercus prinoides</i>); dry seed heads of
+grass (<i>Bromus inermis</i> and <i>B. japonicus</i>).</p>
+
+<p>December 22: Many twigs of bittersweet (<i>Celastrus
+scandens</i>) with fruits still attached; several seed heads of
+sunflower (<i>Helianthus annuus</i>); a few acorns of chinquapin
+oak; fragments of the fruit of osage orange; cured bundles
+of trefoil (<i>Desmodium glutinosum</i>), carrion-flower, and
+tickle grass (<i>Panicum capillare</i>).</p></div>
+
+<p>Although the eastern woodrat is relatively unspecialized in its<span class='pagenum'><a name="Page_525" id="Page_525">[Pg 525]</a></span> feeding
+habits, a few species of favored food plants probably make up the
+greater part of its diet. In northeastern Kansas, at present, osage
+orange probably is by far the most important single species. Despite the
+fact that its aromatic leaves and fruits are somewhat repellent to
+insects and some other animals, they are well liked by woodrats, and
+provide a year-round food supply to those individuals having houses in
+or near the trees. Honey locust similarly provides thorny shelter for
+house sites, while the foliage, the seeds, and the bark of twigs and
+trunks are eaten. In houses that are situated near honey locusts, the
+large, heavy seed pods are sometimes stored by the hundreds. Old pods
+are often used in substitution for sticks as building material in the
+house. Nevertheless, honey locust is used relatively little as compared
+with osage orange. Other plants that figure most importantly in the diet
+include bittersweet, fox grape, pokeberry and horse nettle (<i>Solanum
+carolinense</i>).</p>
+
+<p>Rainey (<i>op. cit.</i>) mentioned that captive woodrats would eat meat, both
+cooked and raw, and on one occasion he found remains of a cicada on a
+house under circumstances suggesting that this insect had been eaten by
+a rat. In the course of trapping for opossums and small carnivores,
+woodrats were caught on many occasions by Fitch in traps baited with
+animal material exclusively&mdash;miscellaneous meat scraps, canned dog-food,
+bacon grease, or carcasses of small vertebrates. In fact, such baits
+seemed to be even more attractive than the grain, seeds, peanut butter
+and raisins that had been used customarily to bait the traps set for
+woodrats. However, such meat baits could be used effectively only in
+cold weather, because of rapid spoilage and interference by insects at
+higher temperatures.</p>
+
+<p>On one occasion an adult pilot black snake found dead on the road, a
+recent traffic victim, was brought to the Reservation headquarters for
+examination and was left overnight in the garage. On the following
+morning the carcass of the snake was found to have been dragged a short
+distance and gnawed; a quantity of flesh was eaten at an exposed wound
+on the neck. Woodrat tracks were thickly imprinted on the dusty soil
+around the snake. The adult male woodrat that lived in the garage had
+evidently spent much time moving about the carcass and over it, and
+feeding upon it. It seemed remarkable that this individual was not
+deterred from feeding on the snake by an instinctive fear of one of its
+chief natural enemies.</p>
+
+<p>Although the eastern woodrat's food consists mostly of vegetation,<span class='pagenum'><a name="Page_526" id="Page_526">[Pg 526]</a></span> the
+strong tendency noted to feed upon flesh when it is available suggests
+that these rodents may, occasionally at least, prey upon helpless young
+of small vertebrates that are readily available to them. Nestling birds,
+either on the ground or in low trees, and young mice in nests that are
+accessible, might tempt the rat to indulge in predation.</p>
+
+
+<h2><i>Breeding</i></h2>
+
+<p>Reproductive activity continues to some extent throughout the year
+except in late autumn and early winter. Presence of a vaginal orifice
+was used as an indication of sexual activity. In most instances the
+orifice was not indicative of actual oestrus, as it persisted through
+the preceding and following stages of an oestrus cycle. In anoestrus the
+orifice is sealed, the genitalia are reduced in size and the skin in the
+genital region is white. Immature females, and adults during most of the
+winter, are in this quiescent condition. Onset of the breeding season in
+late winter is relatively abrupt, and seemingly is a photoperiodic
+response. Breeding may begin in late January, and most females are in
+breeding condition within the first half of February. In oestrus the
+genitalia are enlarged and discolored and the vaginal orifice is
+prominent and gaping. By February most females born the previous season
+have matured, and breeding involves the entire population, except
+possibly for retarded young and individuals suffering from disease,
+injury or malnutrition. Rainey (1956) recorded an average of 2.3 young
+per litter.</p>
+
+<p>Number of litters normally produced in the course of a season by an
+adult female is unknown, but most mature females examined within the
+period February to September inclusive were in some stage of the
+breeding cycle. It is obvious that the females which are successful in
+rearing their litters produce at least two litters annually, and
+probably some produce three litters. When entire litters are lost at an
+early age, to predation, or other causes, productivity is much
+increased, with perhaps only short intervals between pregnancies.</p>
+
+<p>The smallest female having a vaginal orifice weighed 160 grams, but in
+most instances somewhat larger size is attained before the onset of
+oestrus. Judging from the average growth rate of immature females (Fig.
+3), most probably attain sexual maturity at an age of five to six months
+unless this age is reached in the winter period of sexual quiescence.
+Rainey (<i>op. cit.</i>) found no clear cut instances of young maturing in
+time to breed before their first winter. He<span class='pagenum'><a name="Page_527" id="Page_527">[Pg 527]</a></span> concluded, tentatively,
+that in most instances sexual maturity is not attained until the spring
+of the year following that in which the rat is born. However, the
+evidence was inconclusive because few of the young marked survived to
+maturity. In late summer and early autumn, the latter third of the
+breeding season, newly matured young of the year, born in early spring,
+may be the most productive group. Young conceived at the beginning of
+the breeding season, and born in early March, would normally reach adult
+size and breeding maturity in August. For example, a young female first
+caught on June 15, 1951, weighed only 150 grams, but by August 10 she
+had gained to 220 grams (probably in pregnancy) and had a vaginal
+orifice. Of 35 adult and subadult females examined by Fitch in October,
+eleven had a vaginal orifice, the latest on October 18. Of these eleven
+showing signs of breeding, four at least had not yet produced litters,
+judging from the undeveloped condition of their mammae, and others that
+showed evidence of recent lactation probably included young of the year
+that had bred in August or September. One female gave birth to a litter
+in a trap on the night of October 6, 1950. Of 32 adult and subadult
+females recorded by Fitch in November, all were sexually quiescent, with
+the possible exception of one having a partially open vagina on November
+10. All females taken in December, and most of those taken in January,
+also were sexually quiescent. January 20 was the earliest recorded date
+for a female with a vaginal orifice. Females examined in February mostly
+were perforate and many of them appeared to be in oestrus. One female
+trapped on February 19, 1950, weighed only 140 grams and was still
+imperforate. Another, weighing 200 grams on February 3, 1952, still was
+imperforate, but by February 27 she was perforate and appeared to be in
+oestrus. An adult female that appeared to be in oestrus on February 3,
+1952, was imperforate on February 10.</p>
+
+
+<h2><i>Growth</i></h2>
+
+<p>At birth woodrats weigh approximately 10 grams or a little more. In a
+litter born in captivity and kept by Rainey, the average gain amounted
+to a little more than 1.5 grams per day during the first two months, but
+in the third month it was somewhat less. As this was an unusually large
+litter, of five young, one more than the female's teats could
+accommodate, their growth may have been a little less rapid than in most
+of those under natural conditions. At an age of three months they
+averaged approximately 120 grams. The three males consistently exceeded
+the two females.<span class='pagenum'><a name="Page_528" id="Page_528">[Pg 528]</a></span></p>
+
+<div class="figcenter" style="width: 450px;">
+<img src="images/fig3.jpg" width="450" height="413" alt="Fig. 3. Typical growth curves for male and female
+woodrats; early stages are based on the litter of a captive female,
+later stages on average gains of recaptured juveniles and subadults,
+excluding those that seemed to be stunted. Solid line represents males
+and broken line represents females." title="" />
+<span class="caption">Fig. 3. Typical growth curves for male and female
+woodrats; early stages are based on the litter of a captive female,
+later stages on average gains of recaptured juveniles and subadults,
+excluding those that seemed to be stunted. Solid line represents males
+and broken line represents females.</span>
+</div>
+
+<p>Young weighing less than 100 grams are rarely caught in live-traps. Four
+young, all males, first caught at an average weight of 80 grams, gained
+on the average, 1.39 grams per day over intervals that averaged 44 days.
+Six other young males first caught while in the weight range of 100 to
+149 grams, were recaptured after intervals of 17 to 45 days and they had
+gained, on the average, .92 grams per day. The corresponding figure for
+four young females in the same size range was .71 grams per day. In
+seven young males in the weight range 150 to 250 grams, that were caught
+after intervals averaging 66 days, the gain in weight amounted to .83
+grams per day. In seven females in the range 150 to 199 grams, gains
+averaged only .68 grams per day. Fully grown females that are not
+pregnant weigh, most typically, a little less than 250 grams while fully
+grown adult males average a little more than 300 grams. Growth rate and
+adult weight both<span class='pagenum'><a name="Page_529" id="Page_529">[Pg 529]</a></span> are influenced to a large extent by season and even
+more by individual differences. The underlying causes are obscure in
+most instances, but individual rats that are still short of adult size
+may stop growing for periods of months, and some individuals grow much
+more rapidly than others. One male that weighed 108 grams when he was
+first caught on July 3, 1951, was estimated to have been born in early
+May. He was last captured 152 days later on December 2, 1951, and by
+then his weight was 300 grams, representing an increase of 1.2 grams per
+day. Another male that weighed only 75 grams when he was caught on
+October 8, 1950, may have been less than two months old then. By
+November 21, 1951, at a probable age of 15 months, he weighed 350 grams
+having attained almost the maximum size. Other exceptionally large
+individuals were known to be less than two years old, while those rats
+that survived longest on the study areas did not much exceed average
+adult size. These records seem to show that exceptionally large woodrats
+are usually not those of advanced age, but are individuals which have
+grown most rapidly through fortuitous circumstances, probably depending
+upon both innate and environmental factors.</p>
+
+<p>None of the woodrats handled was excessively fat, nor were any
+emaciated. The habit of keeping on hand stores of food at all seasons
+perhaps obviates the necessity for storing quantities of fat. Seasonal
+trends in weight vary among individuals, and are not wholly consistent
+from year to year. Rainey found that in late autumn and winter, rats
+steadily gain weight reaching a peak in late February or March. However,
+in the winters of 1948-49 and 1949-50, Fitch found that most rats lost
+weight and hardly any, even those that were short of adult size, made
+gains.</p>
+
+<p>The following records of a male born in the spring of 1949 show rapid
+growth and attainment of adult size in his first summer, cessation of
+growth during the winter, and resumption of growth, with attainment of
+near-maximum size the following spring.</p>
+
+
+<div class='center'>
+<table border="0" cellpadding="4" cellspacing="0" summary="">
+<tr><td align='left'>June 16, 1949</td><td align='right'>96 gms.</td></tr>
+<tr><td align='left'>September 26, 1949</td><td align='right'>230 gms.</td></tr>
+<tr><td align='left'>September 27, 1949</td><td align='right'>230 gms.</td></tr>
+<tr><td align='left'>October 18, 1949</td><td align='right'>260 gms.</td></tr>
+<tr><td align='left'>October 27, 1949</td><td align='right'>250 gms.</td></tr>
+<tr><td align='left'>October 29, 1949</td><td align='right'>220 gms.</td></tr>
+<tr><td align='left'>November 8, 1949</td><td align='right'>235 gms.</td></tr>
+<tr><td align='left'>November 15, 1949</td><td align='right'>245 gms.</td></tr>
+<tr><td align='left'>November 24, 1949</td><td align='right'>240 gms.</td></tr>
+<tr><td align='left'>November 26, 1949</td><td align='right'>240 gms.</td></tr>
+<tr><td align='left'>November 30, 1949</td><td align='right'>240 gms.</td></tr>
+<tr><td align='left'>December 20, 1949</td><td align='right'>260 gms.</td></tr>
+<tr><td align='left'>February 18, 1950</td><td align='right'>230 gms.</td></tr>
+<tr><td align='left'>April 5, 1950</td><td align='right'>290 gms.</td></tr>
+<tr><td align='left'>April 7, 1950</td><td align='right'>300 gms.</td></tr>
+<tr><td align='left'>October 7, 1950</td><td align='right'>320 gms.</td></tr>
+<tr><td align='left'>November 29, 1950</td><td align='right'>345 gms.</td></tr>
+<tr><td align='left'>March 23, 1951</td><td align='right'>340 gms.</td></tr>
+</table></div>
+
+<p><span class='pagenum'><a name="Page_530" id="Page_530">[Pg 530]</a></span></p>
+
+<p>Another example, showing winter cessation of growth in a male at even
+smaller size is shown below. This was in the winter of 1950-1951.</p>
+
+
+<div class='center'>
+<table border="0" cellpadding="4" cellspacing="0" summary="">
+<tr><td align='left'>November 9</td><td align='right'>145 gms.</td></tr>
+<tr><td align='left'>November 28</td><td align='right'>175 gms.</td></tr>
+<tr><td align='left'>November 29</td><td align='right'>165 gms.</td></tr>
+<tr><td align='left'>January 10</td><td align='right'>180 gms.</td></tr>
+<tr><td align='left'>January 11</td><td align='right'>175 gms.</td></tr>
+<tr><td align='left'>March 1</td><td align='right'>225 gms.</td></tr>
+<tr><td align='left'>March 23</td><td align='right'>200 gms.</td></tr>
+</table></div>
+
+
+<h2><i>Longevity</i></h2>
+
+<p>The longest span of records for an individual woodrat recorded was 991
+days in a female, already adult when she was first caught on November
+18, 1948. Other relatively long spans of records were: 827 days in a
+male, adult when first caught on March 16, 1952; 754 days in a female,
+also adult when first captured; 649 days in a male first captured as a
+juvenile; 465 days in a male, adult when first captured; 409 days in a
+male, juvenile when first captured; 399 days in a female, juvenile when
+first captured; 395 days in a female, adult when first captured; 390
+days in a female, adult when first captured; 366 days in a male, adult
+when first captured. Of these eleven individuals (six females and five
+males) whose records cover more than a year, eight were already adult
+when first caught. These eleven rats represent only 4.3 per cent of the
+total number captured. Our study was made at a time when populations
+were shrinking and disappearing, and obviously individual spans would
+have been longer if we had been working with a stable population. In
+most instances the spans of our records represent only small parts of
+the life spans of the individuals involved. Nevertheless, our records
+emphasize the potentially greater longevity of the woodrat as contrasted
+with the various smaller rodents living in the same area. Of several
+thousand individuals of the genera <i>Mus</i>, <i>Zapus</i>, <i>Reithrodontomys</i>,
+<i>Peromyscus</i>, <i>Sigmodon</i>, and especially <i>Microtus</i>, none is known to
+have survived so long as two years, and only a few individuals are known
+to have survived so long as one year after being marked.</p>
+
+
+<h2><i>Summary</i></h2>
+
+<p>Plant succession resulting from land use practices created habitat
+conditions especially favorable for woodrats in the late nineteen
+forties in northeastern Kansas, and particularly on the University of
+Kansas Natural History Reservation. With protection from prairie fires,
+woody vegetation had encroached onto areas that were formerly grassland,
+and, later, fencing against livestock permitted dense thickets of
+undergrowth to develop. In this region the woodrat<span class='pagenum'><a name="Page_531" id="Page_531">[Pg 531]</a></span> usually lives in a
+forest habitat, and requires for its house sites places that are
+especially well sheltered, as in matted thickets of undergrowth, root
+tangles exposed along eroded gully banks, hollow stumps or tree trunks,
+bases of thorny trees with multiple trunks for support, thick tops of
+fallen trees, or, especially, rock outcrops with deep crevices.</p>
+
+<p>At the time of their maximum population density in or about 1947,
+woodrats probably averaged several per acre on the woodland parts of the
+Reservation. In the autumn of 1948, 17 were caught on the ten-acre tract
+of woodland that was live-trapped most intensively. By then, however,
+the population had already undergone drastic reduction, as shown by the
+fact that there were many unoccupied and disintegrating houses
+throughout the woodland. While the time and manner of mortality was not
+definitely determined, circumstantial evidence suggests that the
+downward trend began in early March, 1948, when record low temperatures
+and unusually heavy snowfall coincided with the time when parturition
+normally occurs. The rigorous weather conditions then may have been
+injurious, not only to the newborn litters but to the females comprising
+the breeding stock. Nevertheless, the population remained moderately
+high through 1948, but by early spring of 1949 more than three-fourths
+of the adults and subadults present in late autumn had been eliminated.
+Again, unusually severe winter weather seemed to be the underlying
+cause, as in January precipitation was the heaviest on record in 81
+years, with penetrating sleet storms, persistent ice glaze, and
+occasional brief thawing followed by sudden drops to extremely low
+temperature.</p>
+
+<p>After the drastic reduction in the winter of 1948-49, the population did
+not recover. Although no further sudden reductions due to extremes of
+weather were noted, the trend seemed to be one of gradual, progressive
+decline throughout the following period of years. Deterioration of the
+habitat, as the developing forest shaded out undergrowth, and inroads of
+certain predators may have been important in preventing recovery of the
+population. Many kinds of predatory mammals, hawks, owls, and snakes
+probably take woodrats occasionally, but the spotted skunk, long-tailed
+weasel, horned owl, timber rattlesnake and pilot black snake are
+considered to be by far the most important predators because of their
+habits and prey preferences. Few actual records of predation on woodrats
+were obtained because of their scarcity during most of the period
+covered by our study.<span class='pagenum'><a name="Page_532" id="Page_532">[Pg 532]</a></span></p>
+
+<p>Of the animals which share the woodrat's habitat, many small mammals,
+reptiles, amphibians, and invertebrates use its houses and live in a
+somewhat commensal relationship.</p>
+
+<p>Woodrats are somewhat territorial, each defending its house and an
+indefinite surrounding area against intrusion by others. Houses tend to
+be spaced at intervals of at least 40 feet; occasionally they are closer
+together. Most foraging for food is done within 75 feet of the house.
+However, woodrats often wander far beyond the limits of the usual home
+range. On the average, males travel more frequently and more widely than
+females, and the larger and older males travel more than the smaller and
+younger. Search for mates provides the chief motivation for wandering.
+Extent of wandering is controlled to a large degree by availability of
+natural travelways, such as rock ledges, by shelters for temporary
+stopping places, such as old deserted houses, and by population density
+of the rats themselves.</p>
+
+<p>Food of the eastern woodrat consists chiefly of vegetation; many kinds
+of leaves, fruits, and seeds are eaten. For many individuals foliage and
+seeds of the osage orange are the staple; hedge rows and dense trees of
+osage orange provide favorable sites for the houses. Woodrats are
+attracted to meat baits, and have been known to feed on flesh of
+carcasses, even on one of the pilot black snake which is a predator on
+the rat.</p>
+
+<p>Woodrats are born blind, naked, and helpless, at a weight approximately
+four per cent of the adult female's. They gain at a rate of at least 1.5
+grams per day in the first two months. When they have reached a weight
+of 100 grams, the gain averages somewhat less than one gram per day, but
+individual variation is great. Males gain more rapidly than females,
+especially in the later stages of growth, as adult weight is greater by
+approximately one-fourth in the male. Some individuals grow to maximum
+adult size at an age of one year. Unusually large individuals are not
+necessarily those that are unusually old. Longevity is greater in
+woodrats than in most smaller rodents. One female of adult size when
+first trapped was last captured 991 days later when she must have been
+well over three years old, and others are known to have survived more
+than two years even though populations were shrinking so that few of the
+rats were able to survive for their normal life span.</p>
+
+
+
+<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_533" id="Page_533">[Pg 533]</a></span></p>
+<h2><i>Literature Cited</i></h2>
+
+
+<p><span class="smcap">Crabb, W. D.</span></p>
+
+<p>1941. Food habits of the prairie spotted skunk in southeastern Iowa.
+Jour. Mamm., 22:349-364.</p>
+
+
+<p><span class="smcap">Fitch, H. S.</span></p>
+
+<p>1947. Predation by owls in the Sierran foothills of California. Condor,
+49:137-151.</p>
+
+
+<p><span class="smcap">Rainey, D. G.</span></p>
+
+<p>1956. Eastern woodrat, Neotoma floridana: natural history and ecology.
+Univ. Kansas Publ. Mus. Nat. Hist., 8: No. 10, in press.</p>
+
+<h4><i>Transmitted March 12, 1956.</i></h4>
+
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Ecological Observations on the
+Woodrat, Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
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+The Project Gutenberg EBook of Ecological Observations on the Woodrat,
+Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
+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: Ecological Observations on the Woodrat, Neotoma floridana
+
+Author: Henry S. Fitch
+        Dennis G. Rainey
+
+Release Date: August 29, 2010 [EBook #33566]
+
+Language: English
+
+Character set encoding: ASCII
+
+*** START OF THIS PROJECT GUTENBERG EBOOK ECOLOGICAL OBSERVATIONS--WOODRAT ***
+
+
+
+
+Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci
+and the Online Distributed Proofreading Team at
+https://www.pgdp.net.
+
+
+
+
+
+
+
+UNIVERSITY OF KANSAS PUBLICATIONS
+MUSEUM OF NATURAL HISTORY
+
+Volume 8, No. 9, pp. 499-533, 3 figs.
+
+June 12, 1956
+
+Ecological Observations on the Woodrat, Neotoma floridana
+
+BY
+
+HENRY S. FITCH AND DENNIS G. RAINEY
+
+UNIVERSITY OF KANSAS
+LAWRENCE
+1956
+
+UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
+Robert W. Wilson
+
+Volume 8, No. 9, pp. 499-533
+Published June 12, 1956
+
+UNIVERSITY OF KANSAS
+Lawrence, Kansas
+
+PRINTED BY
+FERD VOILAND, JR., STATE PRINTER
+TOPEKA, KANSAS
+1956
+
+
+
+
+ECOLOGICAL OBSERVATIONS ON THE WOODRAT, NEOTOMA FLORIDANA
+
+By
+
+Henry S. Fitch and Dennis G. Rainey
+
+
+_Introduction_
+
+The eastern woodrat exerts important effects on its community associates
+by its use of the vegetation for food, by providing shelter in its stick
+houses for many other small animals, and by providing a food supply for
+certain flesh-eaters. In the course of our observations on this rodent
+on the University of Kansas Natural History Reservation, extending over
+an eight-year period, from February, 1948, to February, 1956, these
+effects have changed greatly as the population of woodrats has
+constantly changed in density, and in extent of the area occupied.
+
+This report is concerned with the population of woodrats on the
+Reservation, the changes that the species has undergone, and the factors
+that have affected it. Our two sets of field data, used as a basis for
+this report, supplement each other and overlap little, either in time or
+space. Fitch's field work which covered approximately the western half
+of the Reservation, was begun in September, 1948, and was pursued most
+intensively in the autumn of 1948 and in 1949, with relatively small
+amounts of data obtained in 1950 and 1951 because of the great reduction
+in numbers of rats. Rainey's field work began in the spring of 1951 and
+was continued through 1954, concentrating on a colony in the extreme
+northwestern corner of the Reservation and on adjacent privately owned
+land. In actual numbers of rats live-trapped and for total number of
+records the two sets of data are comparable. Fitch's field work
+consisted chiefly of live-trapping while Rainey's relied also upon
+various other approaches to the woodrat's ecology. Rainey's findings
+were incorporated originally in a more comprehensive report (1956), from
+which short passages have been extracted that are most pertinent to the
+present discussion. Our combined data represent 258 woodrats (153
+Fitch's and 105 Rainey's) caught a total of 1110 times (660 Rainey's and
+450 Fitch's). Rainey's records pertain, in part, to woodrats outside the
+Reservation but within a few hundred yards, at most, of its boundaries.
+
+
+_Habitat_
+
+In the autumn of 1948 the population of woodrats was far below the level
+it had attained in 1947 or earlier, but the rats were still abundant and
+distributed throughout a variety of habitats. Almost every part of the
+woodland was occupied by at least a sparse population. Also, many rats
+lived beyond the limits of the woodland proper, in such places as
+deserted buildings, thickets, roadside hedges, and tangles of exposed
+tree roots along cut banks of gullies. All these situations are
+characterized by providing abundant cover, a limiting factor for this
+woodrat.
+
+In 1947, when the population of woodrats was especially high, plant
+succession on the wooded parts of the Reservation may have been near the
+optimum stage for the rats. For some 80 years, since the time the land
+was first settled and prairie fires were brought under control, woody
+vegetation has been encroaching into areas that were formerly grassland.
+
+About 1934 the University changed its policy with regard to treatment of
+the tract that was later made the Reservation. Up to that time, most of
+the area had been used as pasture and subjected to heavy grazing, but
+several fields had been fenced and cultivated. Under the new policy the
+hillsides and hilltop edges with open stands of various deciduous trees
+were enclosed with stock fences and protected from grazing. Successional
+trends were greatly altered. Woody vegetation, already favored by
+protection from the prairie fires originally important in the ecology of
+this region underwent further development as a result of protection from
+browsing. Thickets of shrubs and saplings sprang up throughout the
+woodland, forming a dense understory layer beneath the discontinuous
+canopy of the relatively scattered mature trees. The composition and
+density of the undergrowth varied markedly in different parts of the
+woodland. The parts that were formerly most open acquired the most dense
+understory. Blackberry, honey locust, osage orange, and prickly ash
+formed in places thorny tangles almost impenetrable to humans. This
+thicket stage reached its peak in density in the middle to late forties
+coinciding approximately with the time of maximum abundance of the rats.
+In the past eight years, under continued protection from burning,
+cutting and browsing, the forest has developed further; sizable trees 20
+feet or more high and up to eight inches in trunk diameter have grown
+from seedlings during the period of protection. An almost continuous
+canopy of foliage has developed, shading the understory and thinning it
+by killing shrubs and saplings. In those situations where the canopy is
+most dense, as on north slopes having stands of young hickory averaging
+twenty feet high, the understory is now largely lacking, but in other
+situations, particularly on south slopes, the understory thickets are
+still dense. On the whole, however, habitat conditions have become less
+favorable for the woodrat.
+
+Within the woodland the population of woodrats was not evenly
+distributed even at its maximum density; only those situations that
+provided sufficient overhead shelter were occupied by woodrats. The
+hilltop limestone outcrop, which was the refugium of the survivors when
+the population was at low ebb, also supported the greatest concentration
+when the population was high. The number of individuals living along any
+particular stretch of ledge could be determined only by intensive
+live-trapping, whereas residences of individuals could be more readily
+identified in most other situations away from the ledge. Stick houses of
+woodrats are, characteristically, large and dome-shaped in woodland, but
+along the ledges they usually lacked this typical form and consisted of
+a much smaller accumulation of sticks, often merely filling a small
+crevice. Sticks carried into such places where they were partly or
+wholly protected from moisture and sunshine were much less subject to
+decay than those in more open situations, and remained long after the
+rats themselves were gone. Accumulations of droppings in depressions in
+rock surfaces beneath overhanging ledges likewise have lasted for many
+years. The rock outcrop provided a continuous travelway along the
+hilltops, and even parts that were not permanently occupied usually had
+some sign. The following types of situations were found to be especially
+favorable for occupancy: deep crevices beneath overhanging projections
+of the ledge; large flat boulders broken away from the main ledge; thick
+clumps of brush (usually fragrant sumac, _Rhus trilobata_) providing
+shelter and support for the house; logs fallen across the ledge
+providing support and protection for the house structure.
+
+A second outcropping limestone stratum approximately 20 feet below the
+level of the hilltop was just as extensive as the upper outcrop, but it
+was little used by the rats because the exposed rock surface was more
+regular, lacking the jagged cracks and deep fissures of the hilltop
+outcrop; and it lacked the overhanging projections which provided
+overhead shelter for the rats along the upper outcrop. More than ninety
+per cent of the rats that were recorded as associated with the outcrops
+were at the hilltop stratum.
+
+Second in preference to the hilltop outcrop as a house site was the base
+of an osage orange tree in thick woods. This tree occurs throughout the
+woodland of the Reservation, having become established when the leaf
+canopy was more open, and the whole area was subject to grazing, with
+less development of the understory vegetation in the woodland. Houses
+were most often situated in those osage orange trees that had been cut
+one or more times, and had regenerated with spreading growth form, the
+multiple branching stems offering substantial support. Occasionally
+houses were built in crotches from two to six feet above ground.
+
+Blackberry thickets also are favorable locations for houses. These
+thickets grew up mostly in fenced areas from which livestock were
+excluded, but where there was not dense shade--hilltop edges and level
+or gently sloping ground adjacent to creek banks. The houses were
+usually in densest parts of the thickets where they were almost
+inaccessible. Mats of dead canes more or less horizontal, with the live
+canes growing up through them, provided effective overhead protection,
+while the ground beneath was relatively open. Houses built in the
+thickets were so well concealed that they were usually not detected
+until after leaves were shed in autumn. In most cases the blackberry
+thickets were small and well isolated. Houses of the rats were sometimes
+unusually near together suggesting that these thickets provided
+especially favorable habitat conditions.
+
+Hollow trees are often utilized, the accumulation of sticks for the
+house being largely inside the cavity. To be suitable for a house site,
+the snag must have an opening near ground level, and another higher on
+the trunk, providing emergency outlets in two directions. Most of the
+hollow trees utilized were black oaks (_Quercus velutina_).
+
+In 1948 there were many houses in cut tops of trees left from small
+scale lumbering operations a few years earlier. The densely branched
+tops of elms, oaks and hickories had satisfied the requirement for
+support of the house and nearby shelter. The houses built in them were
+in open woodland well separated from otherwise favorable situations. By
+1948 the tops were disintegrating and no longer provided effective
+shelter. The houses built in them were falling into disrepair and were
+not permanently inhabited but were often used temporarily by wandering
+individuals.
+
+Along cut banks of gullies where trees were partly undermined by
+erosion, the exposed, tangled root systems provided sites for occupancy.
+In these situations the accumulations of sticks were small and lacked
+the typical domed shape, consisting essentially of a lining to the
+cavity beneath the roots.
+
+Two small buildings at the Reservation headquarters were accessible to
+woodrats and were utilized off and on throughout much of the period of
+this study, despite the fact that most other sites of occupation away
+from the hilltop outcrops were deserted in the same period. One small
+building used as a laboratory had an enclosed wooden box five feet
+square housing an electric water pump. The interior of this box was
+accessible to the rats from beneath the floor. Litter of sticks and
+stems and various food materials were carried in by the rats. The nest
+thus protected and enclosed was not surrounded by the usual accumulation
+of sticks. An old garage 30 feet from the laboratory building was also
+occupied, sometimes by a different individual. The nest and food stores
+were behind boards propped against the wall.
+
+In October, 1948, live-trapping was begun on a heavily wooded slope
+facing northwest, and a ten-acre area was trapped rather thoroughly in
+the succeeding weeks. Because few traps were then available, this was
+the only area that was well sampled in 1948, although diffuse trapping
+was carried on over some 200 acres. On the ten-acre tract a total of 17
+adult and subadult woodrats were caught, four along the hilltop rock
+outcrop, six along the gully at the bottom of the slope, and seven at
+intermediate levels on the slope. Judging from the many unoccupied
+houses, the population on this tract had been much higher before the
+study was begun. On the basis of this sample it seems that in 1947 a
+population of several hundred woodrats lived on the wooded parts of the
+square mile where the Reservation is located.
+
+
+_Reduction of Population_
+
+The abrupt reduction in the population of woodrats on the Reservation
+cannot be explained conclusively with available data. Probably weather
+played a major part, but other unknown factors must have been important
+also. It is certain that the population of woodrats was high, if not at
+an all-time peak, in 1947. In late February, 1948, when one of us
+(Fitch) first visited the area on a preliminary inspection trip (not
+concerned primarily with woodrats), houses of these rats were found to
+be unusually numerous and those seen seemed to be occupied and well
+repaired. Possibly the population was drastically reduced within the
+next few weeks, as unseasonably cold and stormy weather occurred in
+early March. For the first 12 days of March, 1948, temperature averaged
+20 deg. below that of average March weather, and even colder than the
+average for January or February. A reading of -5 deg.F. on March 11 set a
+new low locally for the month since records were begun in 1869. The
+record low temperatures were accompanied by 12.8 inches of snow. This
+spell of unusually severe weather in early March coincided with the
+period in which first litters of young usually are born, as most females
+breed in early February and the gestation period is in the neighborhood
+of five weeks. That most of these first-litter young may have been
+eliminated by the unfavorable extreme of weather at the most critical
+stage in the life cycle may be readily imagined although definite proof
+is lacking. However, the mortality must have extended beyond newborn
+young. Loss of first litters ordinarily would be compensated for by the
+end of the season, since a female usually breeds more than once in the
+course of a season. In any case, by autumn, when the actual field study
+of woodrats was initiated, many houses were already deserted and in
+disrepair. Although the rats were still moderately abundant, they were,
+seemingly, much below the population peak of the preceding year.
+
+Further drastic reduction of adults and subadults took place in the
+winter of 1948-49. In the course of live-trapping operations from
+mid-October into early December, 51 individuals were caught and marked.
+Chiefly because of unfavorable weather conditions, field work was
+discontinued in mid-December, and live-trapping was not resumed until
+early March. Subsequently, only 12 of the woodrats previously marked
+could be recaptured, and the population had become noticeably sparse.
+Seemingly, more than three-fourths of the population present in late
+autumn had been eliminated in the interval. In January, weather was
+exceptionally severe; on the ninth and tenth the worst sleet storm in
+twelve years occurred. Sleet fell in small granules, while the
+temperature remained several degrees below freezing. Partial thawing on
+January 12, 13 and 14 was followed by a steady drizzling rain on the
+fifteenth. On the following day the temperature dropped to -7 deg.F. Ice
+still remained from the sleet storm, and the slush again froze. On the
+night of January 18, there was one of the worst snow storms on record
+and temperature reached a low of 2 deg.F. Exceptionally low temperatures
+persisted through January 24, with more sleet on January 25. Ice from
+the earlier storm still remained. On January 30, the temperature dropped
+to -7 deg. and a three-inch cover of snow still remained over the coat of
+ice. The month of January, 1949, had the heaviest precipitation in 81
+years (5.09 inches) and a cover of ice remained for at least 21 days.
+There were other sleet storms of lesser proportions on February 2 and
+again on February 21.
+
+Ordinarily sleet would not seriously damage woodrats living in houses in
+woodland habitats and less suitable hedge rows because it usually
+freezes as it falls and coats only the surface of the house. Gradual
+thawing would allow normal runoff without much penetration. Because the
+sleet during the storm described above did not form a glaze as it fell,
+the ice particles penetrated many houses. It has been observed many
+times that captive woodrats refused food that was frozen or were unable
+to eat it. Woodrats in live-traps in winter rapidly weaken unless a
+large supply of food is available. If food supplies became sealed over
+by ice, woodrats would have died by starvation or by falling an easy
+prey to predators. The rats were more accessible to several predators
+than were smaller mammals such as meadow voles which were difficult to
+obtain because of the coating of ice over the fields.
+
+The decimated population surviving into the breeding season of 1949
+failed to make substantial gains. In fact, during the following
+four-year period the general trend of the population over the
+Reservation as a whole seemed to be one of gradual further decline.
+
+In November, 1949, the rats were almost gone from the area of north
+slope and hilltop in oak-hickory-elm woodland where the most intensive
+live-trapping and other field work had been done the previous year. The
+following descriptions of houses remaining on the area at that time give
+some idea of the habitat, and of the course of events correlated with
+the fluctuations in numbers of woodrats.
+
+     No. 1. At the hilltop outcrop, partly on a substrate of
+     limestone boulders, built around an elm of two-foot DBH,
+     which lent support to one side. A hackberry sapling one inch
+     in stem diameter grew through the middle of the house,
+     providing further support. The house was two feet high and
+     six feet in diameter, and was in obvious disrepair, with a
+     hole several inches in diameter in its top. It had been
+     occupied in the autumn of 1948. It was constructed mainly of
+     sticks, ranging in diameter from approximately one inch to
+     straw size. Many of the sticks, from .4 to .5 inches in
+     diameter and one to two feet long, seemingly would have been
+     heavy burdens for a rat, although they were of light-weight
+     wood, sumac and elm. Mixed with the sticks were quantities
+     of dry leaves, bark, and chips of wood, all material
+     appearing old and weathered. This house was in
+     elm-oak-hickory woods 50 feet from a cultivated field on the
+     hilltop to the east and south. To the north and west the
+     escarpment sloped away abruptly. There was a coralberry
+     thicket beneath the trees on the adjacent hilltop.
+
+[Illustration]
+
+FIGURE 1
+
+(A) Map of part of University of Kansas Natural History Reservation,
+showing first-capture sites for all woodrats live-trapped in the autumn
+of 1948. Because of the short time involved and the few traps available,
+much of the area shown was not thoroughly trapped. Woodrats were
+abundant, though much less so than in 1947, as shown by the large number
+of deserted houses.
+
+(B) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1949. Woodrats were
+still moderately abundant, but much below the level of the previous
+year. Triangles indicate those capture sites not sampled in 1948.
+
+(C) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1950. Numbers were
+medium-low, having undergone drastic reduction from the peak level.
+Triangles indicate those capture sites where trapping was not done in
+earlier years.
+
+(D) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1951. The
+population was low, but had not yet reached its lowest ebb.
+
+(E) Map of woodrat study area, same as shown in (A), showing
+first-capture sites for all woodrats live-trapped in 1952, when the
+population had declined to relatively low numbers and disappeared from
+much of its former habitat.
+
+(F) Map of the 590-acre Natural History Reservation, showing the area
+where woodrats were studied.
+
+     No. 2. On gently sloping hilltop edge 15 feet from the
+     outcrop and escarpment, built around a forked walnut sapling
+     having both trunks approximately five inches in diameter.
+     The sapling, coming up through the center of the house at a
+     45 deg. angle, evidently had been bent by the accumulated weight
+     of the debris at an early stage of its growth, many years
+     before. Trees were small in this part of the woods, with a
+     well developed understory thicket of coralberry and sumac.
+     This house approximately one foot high and six feet wide,
+     was constructed mainly of sticks and was similar in
+     composition to No. 1, but appeared considerably older with
+     all the sticks blackened and rotten. In the autumn of 1948
+     this house was used by woodrats, but probably only as a
+     temporary stopping place, because it was already in
+     disrepair then.
+
+     No. 3. At edge of escarpment, 25 feet from No. 2, on a flat
+     boulder approximately six feet long, three feet wide and one
+     foot thick. The decaying and much flattened mass of sticks
+     was mainly on top of the boulder, but also spilled over its
+     edges. Fresh sign was noted at this house in the autumn of
+     1948, but the house was already in disrepair then, and
+     seemingly it was used only as a stopping place.
+
+     No. 4. At the hilltop outcrop where an elm had fallen across
+     it. The decaying log remaining was approximately 12 feet
+     long and 15 inches thick. This log passed diagonally through
+     the house, providing its main support. The house was
+     approximately 39 inches high, its summit extending a little
+     above the level of the top of the outcrop. The house was
+     approximately seven feet wide along the outcrop. This house
+     was somewhat intermediate between the typical dome-shaped
+     stick piles that the rat builds in open situations and the
+     formless accumulations of sticks with which some rats living
+     in deep rock crevices line the entrances. Part of the
+     accumulation was beneath the limestone boulders and
+     outcropping slabs. Approximately half of the material used
+     in the house consisted of sticks and the remainder of pieces
+     of bark and chips of wood, mostly gathered from the fallen
+     elm. This house had shrunken noticeably from decay and
+     settling in the months since it was occupied, in the autumn
+     of 1948. The house was surrounded by a thicket of fragrant
+     sumac, dogwood, and hackberry saplings.
+
+     No. 5. At edge of a protruding boulder one foot thick at the
+     hilltop outcrop of the west facing escarpment, and 100 feet
+     back in the woods from the edge of a corn field, in
+     undergrowth of dogwood, wild currant, and coralberry. The
+     house consisted of a pile of rotten twigs, 3 inches deep and
+     30 inches wide on the upper side of the boulder, and a
+     lining of similar material at the lower edge of the boulder,
+     partly blocking the crevice beneath it. The twigs composing
+     the house were old and rotten. However, a few dry but still
+     green hackberry leaves were stored in the crevice beneath
+     the boulder. In a bare space atop the boulder were several
+     recent woodrat droppings, small and obviously produced by an
+     immature individual, which, perhaps, had recently settled at
+     this old house site.
+
+     No. 6. In hilltop woods, 30 feet from a corner adjoining a
+     pasture and a corn field, at the base of an osage orange
+     tree of one foot DBH, and also over a hollow cottonwood log
+     one foot in diameter and three feet from the osage orange
+     tree. Suspended mats of grape and smilax vines, and the
+     thorny, dead, lower branches of the tree provided additional
+     shelter. The house was composed of sticks and twigs, mostly
+     of osage orange, with spines still present; slabs of bark,
+     wood chips, and dry leaves also made up part of it.
+     Materials on the exterior of the house appeared old and
+     weathered, but the house was conical and solid. Seven fresh
+     corn cobs were on the house or near its base, suggesting
+     that corn from the nearby field had figured importantly in
+     the diet of the occupant. A well beaten path led from the
+     base of the house alongside the log, to a large cottonwood
+     tree 15 feet from the house. This evidence that the house
+     was occupied was verified by live-trapping the occupant.
+     Late in 1948, also, the house was occupied by another
+     individual, but seemingly was deserted for a period of
+     months thereafter.
+
+     No. 7. On upper part of north slope where a hickory seven
+     inches in diameter had fallen across an old sunken log
+     approximately one foot in diameter. The house, composed
+     mainly of hickory twigs 1/4-inch to 1/2-inch in diameter,
+     mixed with bark, wood chips, and leaves, was partly decayed,
+     with no fresh sign and was in a thicket of greenbrier,
+     saplings of hickory and hackberry, and cut tops of
+     hickories. The top was flattened to less than four inches
+     above the level of the supporting hickory log. There were
+     large cavities in the side of the house. When first
+     discovered in the autumn of 1948, this house was occupied by
+     a subadult female rat, but she moved away permanently, and
+     the house had been deserted for approximately a year when
+     these observations were recorded.
+
+     No. 8. In middle of northwest slope, in thick branches of
+     broken top of a black oak. This house had become flattened
+     by decay and settling to form a mound approximately one foot
+     high and five feet in diameter. Only the top protruded
+     through the carpet of dry leaves. Once well protected and
+     partly concealed by the branches and twigs of the oak top,
+     this house was now fully exposed by the disintegration of
+     the top. The house consisted chiefly of oak twigs. In
+     October, 1948, a woodrat was live-trapped at this house, but
+     probably it was a wanderer. The house had then already
+     undergone much deterioration.
+
+
+_Natural Enemies_
+
+Some 56 species of animals that regularly prey on small vertebrates live
+on the Reservation. Many of the larger kinds may take woodrats
+occasionally. Because of size, habitat preferences and the time and
+manner of hunting, five species stand out as the more formidable
+enemies--the horned owl (_Bubo virginianus_), prairie spotted skunk
+(_Spilogale putorius_), long-tailed weasel (_Mustela frenata_), pilot
+black snake (_Elaphe obsoleta_) and timber rattlesnake (_Crotalus
+horridus_).
+
+Throughout the study horned owls were common on the area, but their
+numbers were highest in 1948. Samples of pellet collections have shown
+that the cottontail is the staple food, being represented in almost
+every pellet. Various rodents also are important in the diet, the cotton
+rat, prairie vole, or white-footed mouse being most prominent according
+to the time and place of collection. The woodrat is approximately
+optimum size for prey, and it constitutes one of the most preferred food
+sources. Remains of only two woodrats were found in the pellets
+examined, but at times when the pellets were collected woodrats were so
+scarce that they constituted only an insignificant percentage of the
+biomass of potential prey. On several occasions woodrats in live-traps
+were attacked by horned owls, as shown by the overturned and displaced
+trap and quantities of fine down adhering to them and to nearby objects.
+The horned owl lives in the same habitat as does the woodrat. In other
+regions woodrats are known to figure prominently in the diet of the
+horned owl. At the San Joaquin Experimental Range in California, for
+instance, _N. fuscipes_ was found 240 times, more frequently than any
+other kind of prey, in 654 pellets of the horned owl, and this owl was
+shown to be the one most important natural enemy of the rat, although
+many kinds of carnivores, raptors and snakes also took toll from its
+populations. On the Reservation the population of horned owls has been
+fairly stable from year to year, with roughly one pair to 100 acres of
+woodland. Some territories have been maintained continuously throughout
+the eight-year period of observation, though changing to some extent in
+size, shape and area included. In 1948, when livestock grazed on the
+area, and the ground cover of herbaceous vegetation was relatively
+sparse, cottontails were much less abundant than they were later when
+the vegetation was protected. Small rodents including voles, cottonrats,
+and deer mice, were also less abundant then, and the numerous horned
+owls may have been supported in part by the high population of woodrats.
+
+The spotted skunk may be an even more important enemy of the woodrat,
+although the evidence is circumstantial. No records of these skunks
+preying on woodrats have been found in the literature, nor were any such
+instances recorded by us except for attacks on woodrats confined in
+live-traps. This skunk is a formidable enemy of small and medium-sized
+rodents, as it can climb, dig, and squeeze through small openings. That
+it may prey on rat-sized rodents and may even be a limiting factor to
+their occurrence is well shown by Crabb's (1941:353) studies in Iowa. He
+found that Norway rats (_Rattus norvegicus_) ranked third in frequency
+(cottontail, mostly carrion, ranked first) in the winter food of the
+spotted skunk. Crabb observed that about farmyards and farm buildings
+where the skunks had been eliminated by persistent persecution, rats
+were abundant, but that about others where the skunks were present, the
+rats were scarce or absent. On several occasions he noted that heavy
+populations of rats about farm buildings in summer and autumn nearly
+disappeared in winter if a skunk was in residence.
+
+Sign of spotted skunk was noted frequently on various parts of the
+Reservation, especially along the hilltop ledges which were the best
+woodrat habitat. On several occasions skunks released from live-traps
+took shelter in woodrat houses which appeared to be unoccupied.
+According to a local fur dealer, C. W. Ogle, spotted skunks reached a
+peak of abundance in Douglas County in the winter of 1947-1948, and many
+pelts were brought in for sale then. The concentration of skunks may
+have had detrimental effect on the population of woodrats, especially
+when extremes of weather had already made conditions critical for them,
+as in early March, 1948, and in January, 1949, when snow and sleet made
+their usual food supply unavailable.
+
+The long-tailed weasel is considered to be a potentially important enemy
+of the woodrat. Weasels have been seen on the Reservation on only a few
+occasions, but they may be more numerous than these records would
+indicate. Two were caught at the hilltop outcrop, at different times and
+places, in funnel traps put out to catch snakes. The weasel seems to
+prefer this rocky habitat, which is also favored by the woodrat. Because
+of its ferocity and willingness to attack relatively large prey, and
+because it is an agile climber and able to squeeze through any openings
+large enough to accommodate a woodrat, it would seem to be a formidable
+enemy.
+
+The pilot black snake (_Elaphe obsoleta_) is an important enemy of this
+woodrat on the Reservation and probably throughout the rat's geographic
+range except for the extreme western part. Although this snake occurs in
+every habitat of the Reservation, it has been found most often along
+rock outcrops of wooded hilltop edges in the type of habitat most
+favored by the rat. Most often pilot black snakes have attempted to
+escape into crevices of the outcrop. These snakes are also skillful
+climbers and often have escaped by climbing out of reach along branches
+or even vertical tree trunks. On several occasions these snakes have
+been found on or beside woodrat houses, or have escaped into them. Over
+a seven-year period 143 pilot black snakes have been recorded, 53 of
+which were adults.
+
+On September, 1948, a large pilot black snake found basking on a rock
+ledge, distended by a recent meal, was palped and contained a subadult
+female woodrat. On June 19, 1953, one of us, approaching a live-trap set
+under an overhanging rock ledge, saw a four-foot pilot black snake on
+top of it. The snake struck repeatedly at the rat in this trap, but was
+unable to reach it. At each stroke the rat would dash about the trap
+frantically.
+
+These snakes hunt by stealth, and might catch woodrats by entering their
+nests, or by lying in wait along their runways, but are not quick enough
+to catch them in actual pursuit. Young in the nest would seem to be
+especially susceptible to predation by the pilot black snake. These
+snakes hunt by active prowling, either by night or by day, and much of
+their food consists of the helpless young of birds and mammals found in
+the nests. While only well-grown or adult pilot black snakes would be
+able to swallow an adult woodrat, any but first-year young probably
+would be able to overcome and swallow the small young. The female
+woodrat's habit of dragging the young attached to her teats as she flees
+from the house at any alarm must save many litters from predation by the
+pilot black snake. First litters of young, born in early March, are
+already well grown, and past the age of greatest susceptibility to
+predation before the snakes emerge from hibernation in late April or
+early May.
+
+The timber rattlesnake is another potentially destructive enemy, but on
+the Reservation, and throughout much of its original range it is now
+relatively scarce. The genus _Neotoma_ largely coincides in its over-all
+distribution with the genus _Crotalus_, of the rattlesnakes. For most
+kinds of woodrats, the larger species of rattlesnakes are among the
+chief natural enemies.
+
+The timber rattlesnake has habitat preferences similar to those of the
+eastern woodrat. Of 30 timber rattlesnakes recorded on the Reservation
+over an eight-year period, all but one were at or near hilltop rock
+ledges in woodland. The woodrat is probably one of the most important
+prey species for the timber rattlesnake. Like the woodrat, the
+rattlesnake is mostly nocturnal in its activity. Unlike the pilot black
+snake, it hunts by lying in wait, striking prey which comes within
+range, and waiting for it to die from the venomous bite, rather than by
+active prowling. Therefore, it is probably less of a hazard to young in
+the nest than is the pilot black snake. Even young rattlesnakes too
+small to eat woodrats are potentially dangerous to them, as they may
+strike and kill any that come within range.
+
+
+_Commensals_
+
+Rainey (1956) listed many kinds of small animals that use the houses of
+the eastern woodrat and live in more or less commensal relationships
+with these rodents.
+
+A situation unusually favorable for observing woodrats and their
+associates was discovered on the Reservation where, in July, 1948, two
+old strips of sheet metal, each covering an area of approximately 25
+square feet, were used as shelter by a lactating female with three
+young. This was on a brushy slope just below an old quarry site. A rock
+pile and remains of an old rock wall were nearby. Woodrats had carried
+many sticks back under the metal strips, filling the spaces beneath
+their edges. There was a nest and a system of runways beneath the
+strips. In the following seven years this site was seldom deserted for
+long and was used by a succession of individuals. The strips of metal
+could be easily raised and then lowered into place with little
+disturbance. Because the situation was not entirely natural, the
+findings may not be typical of other rat houses. Animals found over a
+period of years beneath these metal strips include: several dozen each
+of the ring-necked snake (_Diadophis punctatus_), five-lined skink
+(_Eumeces fasciatus_), and ant-eating toad (_Gastrophryne olivacea_);
+several individuals each of cottontail (_Sylvilagus floridanus_),
+white-footed mouse (_Peromyscus leucopus_), short-tailed shrew (_Blarina
+brevicauda_), least shrew (_Cryptotis parva_), American toad (_Bufo
+americanus_), Great Plains skink (_Eumeces obsoletus_), pilot black
+snake (_Elaphe obsoleta_); and one each of bull snake (_Pituophis
+catenifer_), spotted king snake (_Lampropeltis calligaster_), red milk
+snake (_L. triangulum_), and timber rattlesnake (_Crotalus horridus_).
+The snakes which were potential predators on the rats seemed to be
+merely utilizing the shelter in these instances, but they may have been
+lying in wait for prey there.
+
+Among mammals, the cottontail and the white-footed mouse are the most
+persistent users of the woodrat houses, especially those that are no
+longer occupied by the rats. On one occasion five white-footed mice were
+caught simultaneously in a trap set beside a house at the base of an
+osage orange tree. Subsequent trapping showed that this house was no
+longer occupied by a rat, but that the mice lived in it. Occupancy of
+such an old woodrat house by white-footed mice may continue long after
+abandonment of the house by the rat, even after the house has partly
+decayed and settled to a small part of its original volume.
+
+Cottontails often have their forms under the edges of houses, either
+occupied or deserted. These situations offer protection overhead and on
+three sides. Abandoned houses having one or more of the entrance holes
+enlarged, as by predators breaking through the side of the house to gain
+access to the nest, are especially well adapted for occupancy by the
+cottontail. The rabbit may make its form inside the house structure.
+
+The opossum, also, finds the type of shelter that it requires in
+abandoned houses that have had the entrances sufficiently enlarged. On
+various occasions opossums or their remains have been found in such old
+houses, and opossums released from live-traps have been known to seek
+shelter in abandoned woodrat houses.
+
+At the old quarry on the Reservation woodrat sign was especially
+abundant. A wooden bin approximately seven feet square, used to store
+crushed rock before quarrying operations were abandoned, was inhabited
+by one rat. At the base of a rock crusher on the top of a bank a few
+yards from the bin was an accumulation of sticks and other debris
+brought by woodrats. A rock wall at the top of the bank between the
+crusher and the bin had many crevices providing shelter for the rats,
+and projecting rocks were littered with their droppings. In the spring
+of 1949 the bin and rock crusher were removed, but at least one rat
+continued to live in the rock wall. In the summer of 1951 several tons
+of corn ruined in the flood were dumped on the top of the bank above the
+wall. By autumn, Norway rats, either brought in with the corn or
+attracted by it, had taken possession of the wall, evidently displacing
+the woodrats, which were no longer present. Although this Old World
+murid rat is much different from the woodrat in habits, it seemingly can
+compete with it and replace it where habitat conditions are otherwise
+favorable for both.
+
+
+_Movements_
+
+The woodrat is dependent on the stick houses that it constructs for
+shelter. For each individual the house constitutes a home base to which
+it is attached, and about which its movements revolve. The area within
+which routine daily movements are confined constitutes the home range,
+which is variable in size and shape. An individual may, and usually
+does, alter its home range over periods of time. The home range is
+somewhat nebulous because the rat may at any time move far beyond the
+small area to which its activities are largely confined. It may be
+motivated by sexual urge or other voluntary wandering; it may be
+enticed by a food supply or some other specific attraction not available
+near its house; or it may be forcibly displaced by an intruder or may
+abandon in favor of an offspring.
+
+An occupied house normally has several runways radiating from it. These
+are well worn paths, smoothed by use, and cleared of obstructions, and
+the rat tends to keep to them in its foraging expeditions. Usually a
+trail leads to a bush or tree showing evidence of heavy use by the rat.
+Ordinarily such a trail cannot be traced more than 30 feet from the
+house, and it seems that the most concentrated foraging occurs within
+this short radius. Experience in live-trapping has indicated that the
+distance covered by a woodrat in its normal foraging for food is
+ordinarily less than 75 feet in any direction from the house.
+
+Usually the rats can be caught in traps only at their houses or nearby
+places that they frequent, as indicated by their sign. When travelling,
+woodrats make use of overhead cover as much as possible. Storing of food
+seems to be associated with the animal's reluctance to wander far from
+home. When a rat is gathering preferred food for storage the home range
+may be enlarged (or the animal may travel beyond the limits of its
+regular home range). In any case the rat may find it necessary to
+traverse an additional area in order to reach the food source. This may
+involve, in part, extension vertically, as when the rat obtains food
+from trees directly over the house. The home range is thus somewhat
+three-dimensional; both trails and feeding places are often above
+ground. Because of dependency on cover, woodrats do not forage randomly
+in all directions from the house.
+
+Although the house and its immediate environs are defended as a
+territory by the occupant, possession may be soon relinquished. A
+woodrat may shift frequently from one house to another, especially if
+unoccupied houses are readily available. Because woodrats had undergone
+drastic reduction in numbers, as discussed on p. 505, unoccupied houses
+in various stages of disrepair were numerous throughout the woodland in
+1948 and 1949, and the rats that were present then seemed especially
+inclined to wander. Even old houses that are collapsed and
+disintegrating may be used temporarily, or may be taken over and
+repaired. Houses that are in sites exceptionally favorable in that they
+provide food and shelter may be occupied more or less permanently, with
+a succession of woodrats over many generations.
+
+Shifts to new areas are perhaps most often motivated by a search for
+mates. Such shifts are, on the average, longer and more frequent in
+males. Males must range farther in search of females when numbers are
+low. On the other hand, when numbers are high and most of the best sites
+are occupied, newly independent young and displaced adults are forced to
+travel greater distances in search of homes. Some of the larger and more
+powerful males move far greater distances than smaller males. The
+longest distances recorded were mostly for large adult males in breeding
+condition. The average maximum distance between successive points of
+capture for 27 adult males was 345 feet. For 39 females (adults and
+subadults) the corresponding figure was 143 feet. The extremes for males
+were 0 to 1080 feet and for females, 0 to 650 feet. Of the 27 males,
+five moved the maximum distance in a single night. Most of the long
+movements by males did not constitute clear-cut shifts in home range,
+and many returned to their original locations.
+
+The average distance between points of first and last captures for 72
+subadult and adult males was 165 feet. A similar figure for 72 subadult
+and adult females was 133 feet. Of the males 23.7 per cent were at the
+same place at the first and last captures; for females the percentage
+was 36.1. These figures are from the combined data of our trapping
+records, but the trends differed sharply in the two sets of records. In
+Fitch's records, movements averaged longer and difference between the
+sexes was much less: 189 feet for 41 males and 178 feet for 42 females.
+Corresponding figures from Rainey's records were: 141 feet for 31 males
+and 74 feet for 30 females. In Fitch's field work, opportunities to
+record exceptionally long movements obviously were better because the
+trap line encompassed a larger area, approximately half a square mile,
+whereas Rainey's live-trapping was concentrated on relatively small
+areas. The reason for the greater vagility of females in Fitch's records
+is less evident. However, the data were obtained within the period of
+drastic population reduction, at a time when there were numerous empty
+houses throughout the woodland, facilitating travel, and shifts from one
+home range to another where conditions were, temporarily at least, more
+favorable. Rainey found that the females in the small colony in woodland
+where he trapped, moved much less than did those that lived along the
+hilltop outcrop, which provided a natural travel route.
+
+Following are several examples of males and females with long histories
+showing individual variation in frequency and distance of movements.
+
+     _Males_
+
+     (1.) First captured October 14, 1951, and last captured 327
+     days later on September 6, 1952. He was taken 12 times. For
+     the first seven captures (October 14, 1951, to July 15,
+     1952), no movements were recorded. In the following seven
+     days he moved 367 feet. Within the next 21 days he returned
+     to within 114 feet of the site of original capture. Less
+     than one month later he was caught for the last time, at
+     this same site.
+
+     (2.) This large male was captured twelve times over a period
+     of 827 days (March 16, 1952, to June 21, 1954). He tended to
+     wander more than other males and was absent from the
+     trapping area from early 1952 to May 1953. One round trip
+     made in a two-weeks period, amounted to a linear distance of
+     1894 feet if the rat followed natural cover. The return trip
+     of 947 feet was the greatest distance traversed in a single
+     night in any of the woodrats we recorded. Other movements
+     between successive captures were: 722, 397, 356, 293, 253
+     and 144 feet (the latter shift made three different times).
+     Sexual urge probably motivated most of his wandering, since
+     numbers of females were low.
+
+     (3.) For this male the span of records was 143 days, with 18
+     captures. For the first eight recaptures, extending over a
+     period of 39 days, he was still at the original location.
+     Four days later he had moved 120 feet and was visiting a
+     female. A week later he returned. In the following month he
+     was recorded as making two more moves, of 115 feet and 215
+     feet. He was last recorded at the hilltop outcrop.
+
+     (4.) The records of this male extended over 465 days, with
+     13 captures. For the entire period only one movement, of 163
+     feet, was recorded. Twelve of the 13 captures were at the
+     same house.
+
+     (5.) This male was captured 16 times over a span of 130
+     days. After the second capture he moved 144 feet along the
+     outcrop and was caught there for the next 14 times, having
+     developed a "trap habit."
+
+     (6.) This male was in the area 210 days (13 captures) and
+     shifted his range. He was first captured on August 17, 1952,
+     at a house at the rock fence 433 feet from the outcrop.
+     Between this date and October 12, 1952, he moved to the
+     outcrop and established residence in a vacant house. He was
+     recorded as making six more moves, the longest of which was
+     only 40 feet.
+
+     (7.) This male was first caught in June, 1949, as a juvenile
+     probably between two and three months old (weighing 96
+     grams) and hence probably still at the maternal house. In
+     September, grown to adult size, he was caught twice, still
+     at this same place. In October, November, December, and in
+     February, 1950, he was caught 11 times at eight places all
+     within a 90-foot radius of his original location. In April,
+     1950, he was caught at points 550 feet WSW and 700 feet SW.
+     In October he was caught within 150 feet of the original
+     location. In November, 1950, and in March and April, 1951,
+     he was caught four times at a place 900 feet SW from his
+     original location.
+
+     (8.) This subadult male was first caught at the hilltop
+     outcrop on October 4, 1949. Two days later he had moved 160
+     feet north along the outcrop. A month later he had shifted
+     600 feet south; in three more days 1040 feet north. On
+     November 15 he was 105 feet south of the November 8
+     location; on November 16, he had moved 70 feet north. On
+     November 17 he had moved 900 feet back south, but had
+     returned on the 18th to the November 16 location. On
+     November 22, he had again shifted 900 feet south. All
+     capture sites were at the hilltop outcrop.
+
+     (9.) This male was caught as a juvenile (75 grams) on
+     October 8, 1950. On November 9 he had moved 220 feet, from
+     the lower outcrop to the upper, and he was recaptured at or
+     near this same site on November 10, 28 and 29, and on
+     January 11 and February 9, 1951. On November 21, 1951, grown
+     to maximum adult size, he was caught at a new location 1080
+     feet from the original.
+
+     (10.) This male was caught as a subadult twice at the same
+     place on November 30 and December 14. By the following
+     autumn he had shifted to a new location 180 feet south along
+     the outcrop, and he was caught there on September 22 and
+     October 18, 1951, and on January 20 and February 2, 1952.
+
+
+     _Females_
+
+     (11.) This female was captured 27 times over a span of 211
+     days. She moved back and forth considerably between two
+     houses 40 feet apart but made only one substantial movement
+     of 245 feet; at this time she was in breeding condition.
+     Nearly seven months after the first capture she was seen for
+     the last time only 16 feet from the original site of
+     capture. It was assumed she fell prey to spotted skunks
+     which were raiding traps.
+
+     (12.) First captured on March 24, 1951, she remained on the
+     area 105 days in which period she was live-trapped 25 times.
+     Sixty per cent of the total captures were at the same house
+     and the longest movement recorded was only 56 feet. She was
+     last caught in a trap 25 feet from the site of original
+     capture.
+
+     (13.) This young adult remained at her house at the rock
+     fence approximately four months. In this period she was
+     captured 11 times. On March 16, 1952, she had moved 410 feet
+     to a house at the eastern section of outcrop, probably
+     searching for a male. She was never seen again.
+
+     (14.) This subadult female moved from the site of original
+     capture to a house 253 feet away on the same outcrop. She
+     was probably in search of a new home when caught the first
+     time. She was recorded at another house 40 feet away on one
+     occasion.
+
+     (15.) Over a span of 90 days and 15 captures this female was
+     not recorded as making any movement. She was living in one
+     of the woodland houses. Mature males were numerous in the
+     area and she was visited by at least two.
+
+     (16.) This female was also living in the woodland section
+     and was first caught on March 30, 1952, in one of the less
+     favorable houses. She was trapped 17 times over a period of
+     85 days. One movement of 68 feet to a new home site was
+     recorded, but the area of foraging probably did not change.
+     She was caught here four times and then disappeared.
+
+     (17.) This female was first trapped as a subadult on October
+     5, 1948, at a house in brush on the upper part of a north
+     slope. On November 24 she had shifted 590 feet to the bottom
+     of the slope and was living in the recess beneath an
+     undermined honey locust on a gully bank. On November 25 she
+     was caught in a similar situation 100 feet farther east
+     along the gully bank. She was recaptured at the gully on
+     November 26 and 30, December 1, 3, 22, and March 8 and 9,
+     and in all she shifted six times between the two gully-bank
+     dens.
+
+     (18.) This female was first trapped as an adult on November
+     18, 1948, in a gully-bank den. She was recaptured at this
+     same place a year later, on November 18 and 30, 1949. On
+     February 19, 1950, she was caught at a hollow sycamore 650
+     feet farther up the gully, and she was recaptured there on
+     February 25 and April 7, and on June 15, 1951. On August 6,
+     1951, she was caught at a house in a thicket on the gully
+     bank, between the first and second locations and 150 feet
+     from the latter.
+
+     (19.) This female was recorded only twice; on October 15,
+     1948, she was at a hilltop rock outcrop. On July 14, 1950,
+     she had moved 1480 feet and was living in a rock pile at the
+     base of the slope, near the same hollow sycamore where
+     female No. 18 had been caught.
+
+     (20.) This female was first caught as an adult on April 5,
+     1950, at a large boulder of a hillside rock outcrop. On
+     October 7, 1950, she had shifted 110 feet to a house at an
+     osage orange tree on the hilltop rock outcrop. On November 9
+     she was back at the first location and on November 28 she
+     had moved 70 feet south along the hillside outcrop. On
+     January 11 and February 9, 1951, she was back at the
+     original location. On November 9, and 21, 1951, she was
+     again at the site 70 feet south, and was still there at her
+     last capture on February 3, 1952.
+
+Ordinarily each house that is in use harbors only a single woodrat. To a
+greater degree than any other kind of mammal on this area woodrats show
+intraspecific intolerance. On various occasions when captives were
+placed in the same or adjacent cages, they focused their attention on
+each other with evident hostility, the more powerful or aggressive
+individuals attacking or pursuing. Several times the confinement of two
+rats in the same live-trap or cage resulted in the death of the weaker
+individual, and seemingly this is the normal outcome unless the attacked
+rat is able to escape. On various other occasions two or more rats have
+been caught in the same trap simultaneously but in every instance these
+were either: a pair of adults, the female appearing to be in oestrus; a
+lactating female and one or more of her young; or young less than
+half-grown, that were obviously litter mates. Older woodrats, especially
+males, often have their ears torn and punctured from fighting.
+
+Territoriality involves, primarily, defense of the house itself. An
+individual that ventures into an occupied house may be quickly routed by
+the occupant even though the latter is smaller. Chasing has been
+observed occasionally, but it is doubtful whether any individual is able
+consistently to defend the entire area over which it forages. Because
+each rat spends most of its time within the shelter of its house, an
+intruder might venture onto its home range unchallenged and undetected,
+so long as it did not enter the nest cavity.
+
+An adult female was live-trapped on October 14, 1951, beside her house
+at the outcrop. As soon as she was released, she disappeared within the
+house. After approximately two minutes, a soft, high pitched whine was
+heard and immediately another woodrat dashed into view closely followed
+by the female. The chase continued for several seconds in the vicinity
+of the house, but the woodrat being chased soon left the area _via_ the
+outcrop. Probably this intruder had moved into the house in the night
+while the female was in the trap.
+
+On June 17, 1952, an adult male was found in a live-trap set at one of
+the brush pile houses in the woodland area. This house was occupied by
+an adult female. He ran into the house after release, and immediately
+there was a loud squeal. He ran outside and paused under some limbs
+approximately 15 feet from the house, and remained there for 15 minutes
+before clipping off an ironweed 12 inches long, which he carried to the
+house. He did not enter the house but stopped beneath overhanging sticks
+at the edge, eating leaves from the plant. He made another attempt to
+enter the house but loud squeals and rustling followed and he returned
+to the ironweed plant and was still eating when observations were
+halted. In another instance, squeals and rustling indicated that the
+occupant and intruder were in combat.
+
+[Illustration: FIG. 2. Diagram illustrating spacing (due to
+territoriality or intolerance of the rats) in twelve woodrat houses in a
+hedge row extending south from south boundary of the Reservation at the
+middle.]
+
+Although home ranges may overlap to some extent, intraspecific
+intolerance tends to maintain a certain minimum interval between houses.
+The arrangement of twelve houses along a hedge row 1170 feet long is
+diagrammatically represented in Figure 2. The average interval was 78.5
+feet (minimum 42; maximum 171). The habitat was uniform. Home ranges
+probably overlap somewhat, and the spacing is the expression of the need
+for an otherwise unoccupied area in which there is sufficient space to
+live. Because individuals tend to fight whenever they meet, there is
+probably a psychological tendency for sequestration which results in
+spacing of houses and reduces social contact thereby avoiding a
+depletion of energy that would be detrimental to the population. Whereas
+condition of the hedge row determines whether or not it will be
+inhabited by woodrats, length determines the number of occupants. The
+spacing of houses in a hedge row must be attributed to something other
+than restriction of sites because the number of sites available always
+exceeds the number that are in use. Although rock outcrops situated in
+areas of uniform habitat have not been observed to the extent that hedge
+rows have, a similar spacing seems to exist and the sites available for
+houses always exceed the actual number found. This behavior pattern
+limits the number of houses and is probably advantageous to the species
+through preventing overcrowding and possible critical depletion of the
+food supply.
+
+Eleven of the young that weighed 100 grams or less when originally
+captured and were presumably still living at the mothers' houses, were
+recaptured repeatedly over periods of weeks or months, providing a
+limited amount of information regarding dispersal. They followed no
+definite pattern. In seven instances (five males and two females) the
+young stayed on at the house beyond the age when they were completely
+independent of the female. In at least two instances the female was
+known to have moved away while the young remained. One female shifted to
+a house 58 feet from the one where she had reared her litter of two, and
+was accompanied by the young male, while the young female stayed on in
+possession of the maternal house. Two months later this young female was
+caught at a house 90 feet away, and an adult male was in possession of
+her former house. One young male shifted to a house 220 feet from his
+original home and remained there several months, but was recaptured once
+back at the original location. Another male made a series of moves over
+a period of weeks and finally settled in a house 490 feet from his first
+home. One male who stayed in the maternal house all summer, and reached
+adult size there, later moved several times, and was last recorded 900
+feet away. One young female shifted 110 feet. In several instances
+juveniles appeared abruptly in houses known to have been unoccupied
+previously, and some of these houses were in poor repair. These young
+had wandered from their maternal houses, for unknown reasons. On one
+occasion a young woodrat was caught in a mouse trap set in a meadow, a
+habitat into which adult woodrats would scarcely be expected to
+venture.
+
+
+_Feeding_
+
+Rainey (1956) has listed 31 food plants that are used by the woodrat in
+northeastern Kansas. He has emphasized that each rat usually obtains its
+food from plants growing in the immediate vicinity of its house, and
+that individuals thus differ greatly in their feeding, according to the
+local vegetation. Therefore, with a sufficiently large number of
+observations, the list of food plants might be greatly expanded, to
+include most of the local flora, with the exception of the relatively
+few kinds that have developed strongly repellent properties rendering
+them unpalatable to herbivores in general.
+
+At the quarry where one or more woodrats usually lived beneath metal
+strips, as described previously (under the heading of "Commensals"), the
+situation seemed to be especially favorable, despite the fact that the
+metal offered no insulation from extremes of heat in summer and cold in
+winter. Perhaps the rat had an alternative nest among nearby boulders,
+to use when temperature was unendurable beneath the metal.
+
+The rat itself, the stored food, and other details of its home life,
+could be observed with a minimum of disturbance by raising one side of
+the metal strip momentarily, then carefully lowering it into place. The
+following observations made in the summer and autumn of 1948 give some
+idea of the range of food plants stored at any one time and the change
+as the season progresses.
+
+     July 12: Bundles of leaves of carrion-flower (_Smilax
+     herbacea_); 15 green pods of honey locust (_Gleditsia
+     triacanthos_) with seeds eaten out; several green fruits of
+     osage orange (_Maclura pomifera_), and several seeds of
+     coffee-tree (_Gymnocladus dioica_).
+
+     July 24: Bundles of green leaves of osage orange and
+     carrion-flower; many pods of honey locust.
+
+     August 30: Three large clusters of the fruits of pokeberry
+     (_Phytolacca americana_).
+
+     October 20: Many small clusters of grapes (_Vitis vulpina_)
+     judged to weigh perhaps one pound in all; several old pods
+     of coffee-tree and a few berries of dogwood (_Cornus
+     Drummondi_) and of pokeberry; a pile of small acorns of
+     chinquapin oak (_Quercus prinoides_); dry seed heads of
+     grass (_Bromus inermis_ and _B. japonicus_).
+
+     December 22: Many twigs of bittersweet (_Celastrus
+     scandens_) with fruits still attached; several seed heads of
+     sunflower (_Helianthus annuus_); a few acorns of chinquapin
+     oak; fragments of the fruit of osage orange; cured bundles
+     of trefoil (_Desmodium glutinosum_), carrion-flower, and
+     tickle grass (_Panicum capillare_).
+
+Although the eastern woodrat is relatively unspecialized in its feeding
+habits, a few species of favored food plants probably make up the
+greater part of its diet. In northeastern Kansas, at present, osage
+orange probably is by far the most important single species. Despite the
+fact that its aromatic leaves and fruits are somewhat repellent to
+insects and some other animals, they are well liked by woodrats, and
+provide a year-round food supply to those individuals having houses in
+or near the trees. Honey locust similarly provides thorny shelter for
+house sites, while the foliage, the seeds, and the bark of twigs and
+trunks are eaten. In houses that are situated near honey locusts, the
+large, heavy seed pods are sometimes stored by the hundreds. Old pods
+are often used in substitution for sticks as building material in the
+house. Nevertheless, honey locust is used relatively little as compared
+with osage orange. Other plants that figure most importantly in the diet
+include bittersweet, fox grape, pokeberry and horse nettle (_Solanum
+carolinense_).
+
+Rainey (_op. cit._) mentioned that captive woodrats would eat meat, both
+cooked and raw, and on one occasion he found remains of a cicada on a
+house under circumstances suggesting that this insect had been eaten by
+a rat. In the course of trapping for opossums and small carnivores,
+woodrats were caught on many occasions by Fitch in traps baited with
+animal material exclusively--miscellaneous meat scraps, canned dog-food,
+bacon grease, or carcasses of small vertebrates. In fact, such baits
+seemed to be even more attractive than the grain, seeds, peanut butter
+and raisins that had been used customarily to bait the traps set for
+woodrats. However, such meat baits could be used effectively only in
+cold weather, because of rapid spoilage and interference by insects at
+higher temperatures.
+
+On one occasion an adult pilot black snake found dead on the road, a
+recent traffic victim, was brought to the Reservation headquarters for
+examination and was left overnight in the garage. On the following
+morning the carcass of the snake was found to have been dragged a short
+distance and gnawed; a quantity of flesh was eaten at an exposed wound
+on the neck. Woodrat tracks were thickly imprinted on the dusty soil
+around the snake. The adult male woodrat that lived in the garage had
+evidently spent much time moving about the carcass and over it, and
+feeding upon it. It seemed remarkable that this individual was not
+deterred from feeding on the snake by an instinctive fear of one of its
+chief natural enemies.
+
+Although the eastern woodrat's food consists mostly of vegetation, the
+strong tendency noted to feed upon flesh when it is available suggests
+that these rodents may, occasionally at least, prey upon helpless young
+of small vertebrates that are readily available to them. Nestling birds,
+either on the ground or in low trees, and young mice in nests that are
+accessible, might tempt the rat to indulge in predation.
+
+
+_Breeding_
+
+Reproductive activity continues to some extent throughout the year
+except in late autumn and early winter. Presence of a vaginal orifice
+was used as an indication of sexual activity. In most instances the
+orifice was not indicative of actual oestrus, as it persisted through
+the preceding and following stages of an oestrus cycle. In anoestrus the
+orifice is sealed, the genitalia are reduced in size and the skin in the
+genital region is white. Immature females, and adults during most of the
+winter, are in this quiescent condition. Onset of the breeding season in
+late winter is relatively abrupt, and seemingly is a photoperiodic
+response. Breeding may begin in late January, and most females are in
+breeding condition within the first half of February. In oestrus the
+genitalia are enlarged and discolored and the vaginal orifice is
+prominent and gaping. By February most females born the previous season
+have matured, and breeding involves the entire population, except
+possibly for retarded young and individuals suffering from disease,
+injury or malnutrition. Rainey (1956) recorded an average of 2.3 young
+per litter.
+
+Number of litters normally produced in the course of a season by an
+adult female is unknown, but most mature females examined within the
+period February to September inclusive were in some stage of the
+breeding cycle. It is obvious that the females which are successful in
+rearing their litters produce at least two litters annually, and
+probably some produce three litters. When entire litters are lost at an
+early age, to predation, or other causes, productivity is much
+increased, with perhaps only short intervals between pregnancies.
+
+The smallest female having a vaginal orifice weighed 160 grams, but in
+most instances somewhat larger size is attained before the onset of
+oestrus. Judging from the average growth rate of immature females (Fig.
+3), most probably attain sexual maturity at an age of five to six months
+unless this age is reached in the winter period of sexual quiescence.
+Rainey (_op. cit._) found no clear cut instances of young maturing in
+time to breed before their first winter. He concluded, tentatively,
+that in most instances sexual maturity is not attained until the spring
+of the year following that in which the rat is born. However, the
+evidence was inconclusive because few of the young marked survived to
+maturity. In late summer and early autumn, the latter third of the
+breeding season, newly matured young of the year, born in early spring,
+may be the most productive group. Young conceived at the beginning of
+the breeding season, and born in early March, would normally reach adult
+size and breeding maturity in August. For example, a young female first
+caught on June 15, 1951, weighed only 150 grams, but by August 10 she
+had gained to 220 grams (probably in pregnancy) and had a vaginal
+orifice. Of 35 adult and subadult females examined by Fitch in October,
+eleven had a vaginal orifice, the latest on October 18. Of these eleven
+showing signs of breeding, four at least had not yet produced litters,
+judging from the undeveloped condition of their mammae, and others that
+showed evidence of recent lactation probably included young of the year
+that had bred in August or September. One female gave birth to a litter
+in a trap on the night of October 6, 1950. Of 32 adult and subadult
+females recorded by Fitch in November, all were sexually quiescent, with
+the possible exception of one having a partially open vagina on November
+10. All females taken in December, and most of those taken in January,
+also were sexually quiescent. January 20 was the earliest recorded date
+for a female with a vaginal orifice. Females examined in February mostly
+were perforate and many of them appeared to be in oestrus. One female
+trapped on February 19, 1950, weighed only 140 grams and was still
+imperforate. Another, weighing 200 grams on February 3, 1952, still was
+imperforate, but by February 27 she was perforate and appeared to be in
+oestrus. An adult female that appeared to be in oestrus on February 3,
+1952, was imperforate on February 10.
+
+
+_Growth_
+
+At birth woodrats weigh approximately 10 grams or a little more. In a
+litter born in captivity and kept by Rainey, the average gain amounted
+to a little more than 1.5 grams per day during the first two months, but
+in the third month it was somewhat less. As this was an unusually large
+litter, of five young, one more than the female's teats could
+accommodate, their growth may have been a little less rapid than in most
+of those under natural conditions. At an age of three months they
+averaged approximately 120 grams. The three males consistently exceeded
+the two females.
+
+[Illustration: FIG. 3. Typical growth curves for male and female
+woodrats; early stages are based on the litter of a captive female,
+later stages on average gains of recaptured juveniles and subadults,
+excluding those that seemed to be stunted. Solid line represents males
+and broken line represents females.]
+
+Young weighing less than 100 grams are rarely caught in live-traps. Four
+young, all males, first caught at an average weight of 80 grams, gained
+on the average, 1.39 grams per day over intervals that averaged 44 days.
+Six other young males first caught while in the weight range of 100 to
+149 grams, were recaptured after intervals of 17 to 45 days and they had
+gained, on the average, .92 grams per day. The corresponding figure for
+four young females in the same size range was .71 grams per day. In
+seven young males in the weight range 150 to 250 grams, that were caught
+after intervals averaging 66 days, the gain in weight amounted to .83
+grams per day. In seven females in the range 150 to 199 grams, gains
+averaged only .68 grams per day. Fully grown females that are not
+pregnant weigh, most typically, a little less than 250 grams while fully
+grown adult males average a little more than 300 grams. Growth rate and
+adult weight both are influenced to a large extent by season and even
+more by individual differences. The underlying causes are obscure in
+most instances, but individual rats that are still short of adult size
+may stop growing for periods of months, and some individuals grow much
+more rapidly than others. One male that weighed 108 grams when he was
+first caught on July 3, 1951, was estimated to have been born in early
+May. He was last captured 152 days later on December 2, 1951, and by
+then his weight was 300 grams, representing an increase of 1.2 grams per
+day. Another male that weighed only 75 grams when he was caught on
+October 8, 1950, may have been less than two months old then. By
+November 21, 1951, at a probable age of 15 months, he weighed 350 grams
+having attained almost the maximum size. Other exceptionally large
+individuals were known to be less than two years old, while those rats
+that survived longest on the study areas did not much exceed average
+adult size. These records seem to show that exceptionally large woodrats
+are usually not those of advanced age, but are individuals which have
+grown most rapidly through fortuitous circumstances, probably depending
+upon both innate and environmental factors.
+
+None of the woodrats handled was excessively fat, nor were any
+emaciated. The habit of keeping on hand stores of food at all seasons
+perhaps obviates the necessity for storing quantities of fat. Seasonal
+trends in weight vary among individuals, and are not wholly consistent
+from year to year. Rainey found that in late autumn and winter, rats
+steadily gain weight reaching a peak in late February or March. However,
+in the winters of 1948-49 and 1949-50, Fitch found that most rats lost
+weight and hardly any, even those that were short of adult size, made
+gains.
+
+The following records of a male born in the spring of 1949 show rapid
+growth and attainment of adult size in his first summer, cessation of
+growth during the winter, and resumption of growth, with attainment of
+near-maximum size the following spring.
+
+    June 16, 1949           96 gms.
+    September 26, 1949     230 gms.
+    September 27, 1949     230 gms.
+    October 18, 1949       260 gms.
+    October 27, 1949       250 gms.
+    October 29, 1949       220 gms.
+    November 8, 1949       235 gms.
+    November 15, 1949      245 gms.
+    November 24, 1949      240 gms.
+    November 26, 1949      240 gms.
+    November 30, 1949      240 gms.
+    December 20, 1949      260 gms.
+    February 18, 1950      230 gms.
+    April 5, 1950          290 gms.
+    April 7, 1950          300 gms.
+    October 7, 1950        320 gms.
+    November 29, 1950      345 gms.
+    March 23, 1951         340 gms.
+
+Another example, showing winter cessation of growth in a male at even
+smaller size is shown below. This was in the winter of 1950-1951.
+
+    November 9             145 gms.
+    November 28            175 gms.
+    November 29            165 gms.
+    January 10             180 gms.
+    January 11             175 gms.
+    March 1                225 gms.
+    March 23               200 gms.
+
+
+_Longevity_
+
+The longest span of records for an individual woodrat recorded was 991
+days in a female, already adult when she was first caught on November
+18, 1948. Other relatively long spans of records were: 827 days in a
+male, adult when first caught on March 16, 1952; 754 days in a female,
+also adult when first captured; 649 days in a male first captured as a
+juvenile; 465 days in a male, adult when first captured; 409 days in a
+male, juvenile when first captured; 399 days in a female, juvenile when
+first captured; 395 days in a female, adult when first captured; 390
+days in a female, adult when first captured; 366 days in a male, adult
+when first captured. Of these eleven individuals (six females and five
+males) whose records cover more than a year, eight were already adult
+when first caught. These eleven rats represent only 4.3 per cent of the
+total number captured. Our study was made at a time when populations
+were shrinking and disappearing, and obviously individual spans would
+have been longer if we had been working with a stable population. In
+most instances the spans of our records represent only small parts of
+the life spans of the individuals involved. Nevertheless, our records
+emphasize the potentially greater longevity of the woodrat as contrasted
+with the various smaller rodents living in the same area. Of several
+thousand individuals of the genera _Mus_, _Zapus_, _Reithrodontomys_,
+_Peromyscus_, _Sigmodon_, and especially _Microtus_, none is known to
+have survived so long as two years, and only a few individuals are known
+to have survived so long as one year after being marked.
+
+
+_Summary_
+
+Plant succession resulting from land use practices created habitat
+conditions especially favorable for woodrats in the late nineteen
+forties in northeastern Kansas, and particularly on the University of
+Kansas Natural History Reservation. With protection from prairie fires,
+woody vegetation had encroached onto areas that were formerly grassland,
+and, later, fencing against livestock permitted dense thickets of
+undergrowth to develop. In this region the woodrat usually lives in a
+forest habitat, and requires for its house sites places that are
+especially well sheltered, as in matted thickets of undergrowth, root
+tangles exposed along eroded gully banks, hollow stumps or tree trunks,
+bases of thorny trees with multiple trunks for support, thick tops of
+fallen trees, or, especially, rock outcrops with deep crevices.
+
+At the time of their maximum population density in or about 1947,
+woodrats probably averaged several per acre on the woodland parts of the
+Reservation. In the autumn of 1948, 17 were caught on the ten-acre tract
+of woodland that was live-trapped most intensively. By then, however,
+the population had already undergone drastic reduction, as shown by the
+fact that there were many unoccupied and disintegrating houses
+throughout the woodland. While the time and manner of mortality was not
+definitely determined, circumstantial evidence suggests that the
+downward trend began in early March, 1948, when record low temperatures
+and unusually heavy snowfall coincided with the time when parturition
+normally occurs. The rigorous weather conditions then may have been
+injurious, not only to the newborn litters but to the females comprising
+the breeding stock. Nevertheless, the population remained moderately
+high through 1948, but by early spring of 1949 more than three-fourths
+of the adults and subadults present in late autumn had been eliminated.
+Again, unusually severe winter weather seemed to be the underlying
+cause, as in January precipitation was the heaviest on record in 81
+years, with penetrating sleet storms, persistent ice glaze, and
+occasional brief thawing followed by sudden drops to extremely low
+temperature.
+
+After the drastic reduction in the winter of 1948-49, the population did
+not recover. Although no further sudden reductions due to extremes of
+weather were noted, the trend seemed to be one of gradual, progressive
+decline throughout the following period of years. Deterioration of the
+habitat, as the developing forest shaded out undergrowth, and inroads of
+certain predators may have been important in preventing recovery of the
+population. Many kinds of predatory mammals, hawks, owls, and snakes
+probably take woodrats occasionally, but the spotted skunk, long-tailed
+weasel, horned owl, timber rattlesnake and pilot black snake are
+considered to be by far the most important predators because of their
+habits and prey preferences. Few actual records of predation on woodrats
+were obtained because of their scarcity during most of the period
+covered by our study.
+
+Of the animals which share the woodrat's habitat, many small mammals,
+reptiles, amphibians, and invertebrates use its houses and live in a
+somewhat commensal relationship.
+
+Woodrats are somewhat territorial, each defending its house and an
+indefinite surrounding area against intrusion by others. Houses tend to
+be spaced at intervals of at least 40 feet; occasionally they are closer
+together. Most foraging for food is done within 75 feet of the house.
+However, woodrats often wander far beyond the limits of the usual home
+range. On the average, males travel more frequently and more widely than
+females, and the larger and older males travel more than the smaller and
+younger. Search for mates provides the chief motivation for wandering.
+Extent of wandering is controlled to a large degree by availability of
+natural travelways, such as rock ledges, by shelters for temporary
+stopping places, such as old deserted houses, and by population density
+of the rats themselves.
+
+Food of the eastern woodrat consists chiefly of vegetation; many kinds
+of leaves, fruits, and seeds are eaten. For many individuals foliage and
+seeds of the osage orange are the staple; hedge rows and dense trees of
+osage orange provide favorable sites for the houses. Woodrats are
+attracted to meat baits, and have been known to feed on flesh of
+carcasses, even on one of the pilot black snake which is a predator on
+the rat.
+
+Woodrats are born blind, naked, and helpless, at a weight approximately
+four per cent of the adult female's. They gain at a rate of at least 1.5
+grams per day in the first two months. When they have reached a weight
+of 100 grams, the gain averages somewhat less than one gram per day, but
+individual variation is great. Males gain more rapidly than females,
+especially in the later stages of growth, as adult weight is greater by
+approximately one-fourth in the male. Some individuals grow to maximum
+adult size at an age of one year. Unusually large individuals are not
+necessarily those that are unusually old. Longevity is greater in
+woodrats than in most smaller rodents. One female of adult size when
+first trapped was last captured 991 days later when she must have been
+well over three years old, and others are known to have survived more
+than two years even though populations were shrinking so that few of the
+rats were able to survive for their normal life span.
+
+
+
+
+_Literature Cited_
+
+
+CRABB, W. D.
+
+1941. Food habits of the prairie spotted skunk in southeastern Iowa.
+Jour. Mamm., 22:349-364.
+
+
+FITCH, H. S.
+
+1947. Predation by owls in the Sierran foothills of California. Condor,
+49:137-151.
+
+
+RAINEY, D. G.
+
+1956. Eastern woodrat, Neotoma floridana: natural history and ecology.
+Univ. Kansas Publ. Mus. Nat. Hist., 8: No. 10, in press.
+
+_Transmitted March 12, 1956._
+
+
+
+
+
+
+End of the Project Gutenberg EBook of Ecological Observations on the
+Woodrat, Neotoma floridana, by Henry S. Fitch and Dennis G. Rainey
+
+*** END OF THIS PROJECT GUTENBERG EBOOK ECOLOGICAL OBSERVATIONS--WOODRAT ***
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