Comparative Ecology of Pinyon Mice and Deer Mice in Mesa Verde National Park Part 5

MICROCLIMATES OF DIFFERENT HABITATS

Four microclimatic stations were established in units D, F, L and M of the trapping grid to record air temperatures and relative humidities at ground level. These sites were chosen as being representative of larger topographic or vegetational areas within the grid. Belfort hygrothermographs were installed on June 10, 1964, and were serviced once each week through October 31, 1964, at which time the stations were dismantled. Each station consisted of a shelter 18 by 9 by 11.5 inches, having a false top to minimize heating (Fig. 9). The shelters were painted white. Several rows of holes, each one inch in diameter, were drilled in all four sides of each shelter, to provide circulation of air. The holes were covered by bra.s.s window screening to prevent entry of insects and rodents. Preliminary tests with several U. S. Weather Bureau maximum and minimum thermometers, suspended one above the other, from the top to the bottom of the shelter, revealed that there was no stratification of air within the shelters. Nevertheless, each shelter was placed so that the sun did not strike the sensing elements of the hygrothermograph inside it.

[Ill.u.s.tration: FIG. 9: (above) Photograph of microclimatic shelter built to house hygrothermograph. False top minimizes heating, and ventilation holes are covered with screening. (below) Photograph showing shelter in use.]

Accuracy of the hair elements was checked by means of a Bendix-Friez battery driven psychrometer, in periods when humidity conditions were stable (on clear days the relative humidity is at its lowest limits and is "stable" for several hours during early afternoon).

The four microclimatic stations were in the following places: 1) a stand of big sagebrush near Far View Ruins; 2) a pinyon-juniper-muttongra.s.s a.s.sociation; 3) a stand of big sagebrush at the head of a drainage; and 4) a stand of Gambel oak on a southwest-facing slope of the drainage.

Table 4 shows monthly averages of maximum and minimum air temperatures and relative humidities at each of the four sites. Vegetation and microclimates of the sites are discussed below.

_Far View Sagebrush Site, 7,650 feet elevation_

The shelter housing the hygrothermograph was next to the stake of station F4a in the trapping grid (Fig. 10), in a stand of big sagebrush on the flat, middle part of the mesa top, approximately 100 yards southwest of Far View Ruins. The sagebrush extends approximately 200 feet in all directions from the station (Fig. 5). Pinyon pine and Utah juniper trees are encroaching upon this area, and scattered trees are present throughout the sagebrush. This area is one of the habitats of _P. maniculatus_.

Sagebrush tends to provide less shade for the ground than pinyon-juniper woodland, and therefore the surface temperatures of the soil rise rapidly to their daily maximum. In mid-June, air temperatures rise rapidly from 6 A. M. until they reach the daily maximum between 2 and 4 P. M. Shortly after 4 P. M. the air temperatures decrease rapidly and reach the daily low by about 5 A. M.

Relative humidities follow an inverse relationship to air temperatures; when air temperatures are highest, relative humidities approach their lowest values. Thus, on clear days, humidities decrease during the day, reaching a minimum slightly later than air temperatures attain their maximum. Unless it rains, the highest humidities of the day occur between midnight and 6 A. M.

_Drainage Site, 7,625 feet elevation_

This site was in the bottom of the drainage that runs through the eastern side of the trapping grid, and through parts of units M, N, I, and J. The site was at station M4d on a level bench at the head of the drainage (Fig. 11). Southward from the station the drainage deepens rapidly, and the bottom loses approximately 25 feet in elevation for every 200 feet of linear distance. _P. maniculatus_ lives here.

The microclimate of the drainage differs markedly from that of other stations. The major difference is attributable to the topography of the drainage itself. Nocturnal cold air flows from the surrounding mesa top to lower elevations. A lake of cold air forms in the bottom of the drainage; the depth of the lake depends in part upon the depth of the drainage. The same phenomenon occurs in canyons and causes cooler night time temperatures on the floor of canyons than on adjacent mesa tops (Erdman, Douglas, and Marr, in press). Drainage of cold air into lower elevations affects both nocturnal air temperatures and relative humidities. Table 4 shows that maximum air temperatures in the drainage did not differ appreciably from those at other stations. Mean minimum temperatures, however, were considerably lower in the drainage than at the other sites. This phenomenon is reflected also in the mean air temperatures at this station.

[Ill.u.s.tration: FIG. 10: (above) Photograph of microclimatic station at the Far View Sagebrush Site, at trapping station F4a in the grid south of Far View Ruins. Dominant vegetation is _Artemisia tridentata_.]

[Ill.u.s.tration: FIG. 11: (below) Photograph of microclimatic station at the Drainage Site, in the bottom of a shallow drainage at trapping station M4d of the grid south of Far View Ruins.]

The drainage site had the highest humidities of all stations each month in which data were collected (Table 4). Relative humidities of 90 to 100 per cent were common in the drainage, but occurred at other stations only in rainy periods. For example, in the month of August, 26 of the daily maximum readings were between 95 and 100 per cent at the drainage site, but at the other stations relative humidities were above 95 per cent for an average of only nine nights. Minimum humidities were about the same for all stations, since they are affected by insolation received during the day, and not by the drainage of cold air at night.

_Oak Brush Site, 7,640 feet elevation_

The station was in an oak thicket at trapping station L4a, 250 feet south and 50 feet east of the drainage site on a southwest-facing slope of about 30 degrees (Fig. 12). The station was on the lower third of the slope, approximately 15 feet higher than M4d, the station in the bottom of the drainage. _P. truei_ and _P. maniculatus_ occur together in this area.

Air temperatures and relative humidities at this station did not differ appreciably from mean temperatures and humidities at the other stations.

The unusual feature is the lack of evidence of cold air drainage. The lake of cold air in the bottom of the drainage apparently is too shallow to reach this station. This site is near the head of the drainage, and the cold, nocturnal air probably moves rapidly down slope into the deeper parts of the canyon, rather than piling up at the shallow head of the drainage.

In spite of the shade afforded the ground by the oak brush, temperatures reached the same maximum values as at the drainage site, owing to the orientation of the slope. South-facing slopes receive more direct insolation throughout the day and throughout the year than north-facing slopes and mesa tops (Geiger, 1965:374). In Mesa Verde, south-facing slopes tend to be more arid; snow melts rapidly, and most of this moisture evaporates. As a consequence, south-facing slopes have less soil moisture and more widely-distributed vegetation than north-facing slopes where snows often persist all winter and melt in spring. (For a detailed discussion of climates on northeast-versus-southwest-facing slopes in Mesa Verde, see Erdman, Douglas, and Marr, in press.)

_Pinyon-Juniper-Muttongra.s.s Site, 7,600 feet elevation_

The station was in the trapping grid at D5b (Fig. 13). The pinyon-juniper woodland surrounding this site resembles much of the woodland on the middle part of the mesa. The forest floor is well shaded by the coniferous canopy, and muttongra.s.s is the dominant plant in the ground cover. _P. truei_ lives in this habitat.

The climate at this site is moderate. Shade from the canopy greatly moderates the maximum air temperatures during the day; minimum air temperatures, however, are about the same as at the other stations (Table 4). Mean temperatures are somewhat lower at this site than at the others because of the lower maximum temperatures. Relative humidities do not differ markedly from those at other stations.

Figure 14 shows hygrothermograph traces at all stations for a typical week. An interesting phenomenon is ill.u.s.trated by several of these traces. By about midnight, air temperatures have cooled to within a few degrees of their nightly low. At this time, heat is given up by the surface of the ground in sufficient quant.i.ties to elevate the air temperature at ground level. This release of reradiated energy lasts from one to several hours, then air temperatures drop to the nightly low just before sunrise. A depression in the percentage of relative humidity accompanies this surge of warmer air. On some nights winds apparently disturb, or mix, the layers of air at ground level. On such nights the reradiation of energy is not apparent in the traces of the thermographs.

Reradiation of energy is restricted to ground level, and traces of hygrothermographs in standard Weather Bureau shelters, approximately four feet above the ground surface, at other sites on the mesa top did not record it.

[Ill.u.s.tration: FIG. 12: (left) Photograph of microclimatic station at the Oak Brush Site, at trapping station L4a of the grid south of Far View Ruins. (right) General view of the stand of Gambel oak in unit L of the trapping grid.]

[Ill.u.s.tration: FIG. 13: Photograph of microclimatic station at the Pinyon-Juniper-Muttongra.s.s Site, at trapping station D5b of the grid south of Far View Ruins. Gra.s.s in the foreground is muttongra.s.s, _Poa fendleriana_.]

The instruments used in this study were unmodified Belfort hygrothermographs containing as sensing units a hair element for relative humidity and a Bourdon tube for air temperatures. The hair element, especially, does not register changes in humidity at precisely ground level; rather, it reflects changes in the layer of air from about ground level to about a foot above. Thus data from these instruments give only approximations of the conditions under which mice live while they are on the ground.

Climatic conditions greatly influence trapping success. Larger numbers of mice generally were caught on nights when humidities were higher than average. Rain in part of the evening almost invariably resulted in more mice of each species being caught. This was probably due to increased metabolism, by the mice, to keep warm. Apparently the mice began foraging as soon as the rains subsided; mice were always dry when caught after a rain. Few mice were caught if rains continued throughout the night and into the daylight hours.

TABLE 4--Monthly Averages of Daily Means for Maximum, Minimum, and Mean Air Temperatures and Relative Humidities at Four Sites in Mesa Verde National Park, Colorado.

===================+========================+======================== Site Maximum Temps. Maximum R. H.

J J A S O J J A S O Far View Sagebrush 89 91 86 77 74 68 84 82 88 71 Drainage 86 91 85 78 78 87 94 93 96 84 Oak Brush 86 88 82 76 81 57 78 80 80 66 Pinyon-Juniper-Poa 75 80 74 66 64 59 83 82 88 58 Minimum Temps. Minimum R. H.

J J A S O J J A S O Far View Sagebrush 42 53 50 42 31 18 24 25 29 21 Drainage 36 48 45 38 26 21 26 27 29 30 Oak Brush 42 52 50 42 32 19 25 30 31 21 Pinyon-Juniper-Poa 44 54 50 42 34 22 30 29 32 25 Mean Temps. Mean R. H.

J J A S O J J A S O Far View Sagebrush 66 72 68 60 52 43 54 54 48 46 Drainage 61 70 65 58 52 54 60 60 62 52 Oak Brush 64 70 66 59 56 38 51 55 56 44 Pinyon-Juniper-Poa 60 67 62 54 49 41 56 55 60 42 -------------------+------------------------+------------------------

[Ill.u.s.tration: FIG. 14: Diagram of hygrothermograph traces showing daily progressions of air temperatures and relative humidities at each of four microclimatic stations, from the morning of July 1 through the morning of July 8, 1964. Slanting vertical lines on each chart designate midnight (2400 Hrs.) of each day.]

Nights of high trapping success usually were a.s.sociated with days having solar insolation below the average. Insolation was measured with a recording pyrheliometer at a regional weather station (M-2) on the middle of Chapin Mesa, at an elevation of 7,150 feet (Erdman, Douglas, and Marr, in press). This station was approximately one mile south of the trapping grid; isolation at this site would have been essentially the same as that received by the trapping grid. Below-average isolation for one day indicates cloudy conditions, which are accompanied by increased humidity, but may or may not be accompanied by precipitation.

Trapping on nights preceded and followed by days of average or above average isolation with average humidities--indicative of clear days and clear moonlit nights--did not yield appreciably higher catches of mice than other nights. Hence there was no evidence that mice tended to avoid, or to seek out, traps on clear moonlit nights.

On cold, humid nights in autumn numerous mice caught in Sherman live traps succ.u.mbed from exposure, even though nesting material (kapok or cotton) and food were in the traps. Occasionally mice succ.u.mbed to heat when traps were inadvertently exposed to too much sunlight. Apparently little heat is required to kill individuals of either species. Traps in which animals died due to excessive heat usually were not hot to the touch; in most instances the traps were checked before 9:00 A. M., several hours before the sun caused maximum heating. Such individuals may have licked the fur of their chests in an attempt to lower their body temperatures. Although mice characteristically salivate before succ.u.mbing from heat, these individuals had moist fur over the entire chest and upper parts of the front legs, indicating licking. Mice killed by exposure to heat or cold usually were juveniles or young; subadult and adult individuals of both species were more tolerant. Older animals would be expected to have better homeostatic controls than younger individuals.

HABITAT PREFERENCE

In Mesa Verde _P. truei_ and _P. maniculatus_ occur together only at the fringes of the pinyon-juniper woodland, where ecotonal areas provide less than optimum habitats for both species. Almost all individuals of _P. truei_ occur only in pinyon-juniper woodland, whereas _P.

maniculatus_ occurs only in more open habitats, such as gra.s.sy meadows and stands of sagebrush.

Pinyon mice were abundant in a variety of a.s.sociations within the pinyon-juniper woodland. The highest population densities were in pinyon-juniper woodland having an understory of mixed shrubs. In such an a.s.sociation, _Poa fendleriana_ usually is the dominant gra.s.s in the ground cover. _P. truei_ was especially abundant along brushy slopes where mixed shrubs (_Amelanchier_, _Cercocarpos_ and _Fendlera_) were codominant with pinyon pines and Utah junipers. The pinyon-juniper-mixed shrub area west of Far View Ruins was almost optimum habitat for _P.

truei_.

_P. truei_ was abundant on the rocky ridge of Wetherill Mesa near Mug House; the pinyon-juniper woodland here has a _Cercocarpos_ understory, and appears to provide close to optimum conditions for this species.

Not all a.s.sociations of the pinyon-juniper woodland support large numbers of _P. truei_. Pinyon-juniper woodland having a ground cover of _Poa fendleriana_, and no shrubs, supports few mice; the woodland on Wetherill Mesa near Long House is an example. Juniper-pinyon woodland having a _Purshia tridentata_ understory also supports only a few mice.

Such areas occur on the southern ends of the mesas and are characterized by widely-s.p.a.ced trees and little ground cover--a reflection of the relatively low amounts of precipitation received by the southern end of the park.

_P. truei_ was not found in gra.s.slands on Navajo Hill, or in meadows at the southern end of Moccasin Mesa. The old burned areas on the northern end of Wetherill Mesa and on Morfield Ridge now support numerous gra.s.ses and shrubs, but _P. truei_ appears not to live there.

_P. truei_ tends to avoid stands of sagebrush, or gra.s.slands, lacking pinyon or juniper trees. _P. truei_ may venture into such areas while feeding. This species is found in thickets of Gambel oak and in areas with an overstory of mixed shrubs only when a living pinyon-juniper canopy is present, or when a woodland adjoins these areas.

Rocky terrain apparently is not a requirement for _P. truei_, since much of the pinyon-juniper woodland that is free of rocks supports large numbers. Optimum habitat, however, had a rocky floor. In such places, rocks probably are of secondary importance, whereas the shrubs and other plants growing on rocky soils are important for food and cover. Rocks likely provide additional nesting sites, and allow a larger population to live in an area than might otherwise be possible.