Temperature Relationships

In the locality of my study racers spend approximately half the year

in winter dormancy. Earliest spring records and latest fall records

for 13 years are shown in Table 4. The spring records in nearly all

instances pertain to snakes found in the open or beneath flat rocks

warmed by sunshine, usually at or near the rock ledges where

hibernation occurs. Juveniles are especially well represented in these

earliest spring records, and it seems that they tend to emerge a

little earlier, on the average, than the adults, either because they

have hibernated in more superficial and less well insulated situations

or because their lesser body mass permits more rapid warming to

activity than can occur in the adults. The latest fall records all

pertain to racers trapped along the rock outcrops, and none was a

young of the year.

Table 4. Earliest and Latest Recorded Annual Dates When Blue Racers

Were Active on the Reservation or Rockefeller Tract

==========+==========+=============++==========+==========+============

| Earliest | Latest || | Earliest | Latest

Year | spring | fall || Year | spring | fall

| record | record || | record | record

----------+----------+-------------++----------+----------+------------

1950 | April 16 | November 4 || 1957 | April 30 | November 2

1951 | April 19 | November 13 || 1958 | April 25 | November 20

1952 | April 23 | November 12 || 1959 | April 6 | November 10

1953 | April 8 | November 12 || 1960 | April 22 | November 6

1954 | April 20 | November 12 || 1961 | April 19 | November 1

1955 | April 15 | November 2 || 1962 | April 23 | October 27

1956 | May 11 | November 14 || | |

----------+----------+-------------++----------+----------+------------

Most of the population undoubtedly emerged somewhat later than the

average date of April 16 indicated by the records in Table 4, and

retired somewhat earlier than the average date of November 8. However,

a small percentage of the population probably emerged even earlier

each year than my records indicate, and retired into hibernation later

than my records indicate. In a typical year, temperatures in April and

early May are only occasionally above the level at which racers are

able to become active, but are below this threshold most of the time.

The same statement applies to an autumn period of late October and

November. Most racers are dormant in their hibernacula during these

transitional periods of spring and autumn, but some--those that have

emerged early in spring, or those that have not yet retired (in

fall)--retreat to temporary shelters and revert to a semi-torpid state

when temperatures fall below the critical level.

Fig. 3 shows the relative extent of activity along the hilltop

outcrops, as reflected by numbers of racers caught at different times

during the autumn. Data from 14 years are combined, and the large

composite sample indicates that in an average year there is relatively

little activity along the hilltop outcrops in early September, but

that activity rapidly increases to a peak in mid-October and then

tapers off rapidly, usually ending in mid-November, but occasionally

ending as early as late October or as late as late November.

The racers recorded in traps had, in many instances, been confined in

them for from one to three days before they were found. For any one

year records are not sufficiently numerous to show the trend as well

as Fig. 3, but Fig. 4 shows year-to-year differences; 1958 was a

fairly typical year, and also was the year in which the largest sample

was obtained; in 1949 the largest catches were made earlier than

usual, and the racers retired early into hibernation; in 1954 warm

weather persisted until unusually late in autumn, and racers remained

active beyond the time when they ordinarily would have been

hibernating; in 1955 and 1961 the most concentrated activity along the

outcrops, as reflected by day to day catches, came later than usual,

but unseasonably cold weather ended all activity abruptly, earlier

than usual.

[Illustration: Fig. 3. Records of blue racers trapped along hilltop

limestone outcrops in autumn, a composite sample of 14 years (1949

through 1962) from the Reservation and Rockefeller Tract, showing the

catch grouped in ten-day intervals, beginning with September 1 to 10

and ending with November 20 to 29. Averages of the maxima, means, and

minima of daily temperatures for each period are shown.]

The racer's annual cycle of activity is, of course, controlled

primarily by the weather, and is much delayed or accelerated in some

years. But certain stabilizing factors cause the racer's annual cycle

to be less variable than that of the weather. For example, in spring

when persistently cool weather delays emergence from hibernation

beyond the normal time, the increasing azimuth of the sun, and more

intense sunshine cause the soil to warm, despite low air temperature,

until emergence is finally triggered. Having once emerged, the racer

is able to control its bodily temperature to a large extent by basking

in sunshine to gain warmth, or by seeking shade or underground shelter

to escape overheating. By such behavioral thermoregulation extremes of

weather are neutralized, or at least buffered to some degree.

[Illustration: Fig. 4. Yearly variation in catch of blue racers along

hilltop outcrop in autumn on the Reservation and Rockefeller Tract,

grouped in same ten-day intervals indicated for Fig. 3.]

In the course of my study no racers were found in their actual

hibernacula. Insofar as known, these were always in deep crevices in

strata of limestone near hilltops, and were inaccessible except by

removal of the solid rock. The crevices where racers hibernate are

known to be several feet deep in some instances, extending well below

the frost line. Periodic temperature readings in two such crevices at

depths of 12 inches and 30 inches, taken in the winter of 1954 (Fitch,

1956:471) showed that temperatures during dormancy are usually well

within the range 0°C to 10°. Whether the racers congregate in

hibernating masses in regular "dens" on the Reservation has not been

definitely determined, but if so, most of the hibernating groups must

be small, because those trapped along the ledges are well scattered,

and, in fact, give the impression of being rather uniformly

distributed along them. However, ledges of northward exposure are not

used as hibernation sites, so far as known, and those of full

southward exposure are perhaps preferred, especially where the hilltop

has a southward projecting spur, and the exposed rock face is

extensive, with many cracks and fissures. I have been unable to detect

differences in types of hibernation sites preferred between the racer

and the copperhead, which is somewhat more numerous on the same area.

Several authors have contributed to knowledge of hibernation in the

racer. Boyer and Heinze in Missouri (1934:195) noted that blue racers

often were associated with copperheads in the vicinity of places

chosen for hibernation. Burt (1935:329) in Kansas found blue racers

emerging from dens among rocks on a prairie hillside, associating with

other snakes, _Diadophis punctatus_, _Elaphe guttata_, and _Pituophis

melanoleucus_. In the Chicago region, Pope (1944:173) reported scores

of blue racers aggregating in October on and around a sand dune with

oak woodland. In Ohio, Conant (1938:55) found three blue racers

hibernating together about three feet underground in a small hole. One

found at another locality had apparently hibernated in company with a

massasauga (_Sistrurus catenatus_). In Maryland Cohen (1939:137) saw

racers (_C. c. constrictor_) in the act of emerging from an old vole

burrow that was a communal hibernaculum, on April 6, 8, and 10. Air

temperatures at the time of emergence were 12.5° and 18.5° Centigrade.

In Illinois, Schroder (1950:1-2) found seven blue racers hibernating

in masses, intertwined with each other and with bull snakes at depths

of 36 inches and 42 inches in an abandoned mammal burrow in a sand

dune area in early February.

[Illustration: Fig. 5. Bodily temperatures of blue racers kept in a

large outdoor enclosure and checked from time to time when they were

active and the sun was shining, in June and July, 1962.]

In the course of routine field work I often carried a Schultheis

quick-reading thermometer, and from time to time I had opportunities

to take the body temperatures of blue racers newly captured by hand.

The trend of these records indicated the temperature range within

which the snakes normally limit their activity, and the preferred

temperature. In an earlier publication (Fitch, 1956:459-460) based on

a few more than half the number of records of temperature now

available, I discussed responses of the blue racer to temperature. The

newer data bear out the trends previously revealed; of 60 records, 39

are within the six-degree range from 29° to 35°, and records are most

concentrated in the one-degree range, 34° to 35°. Racers were found

active at air temperatures between 15.5° and 32.4°, with the greatest

concentration between 26° and 27°. Compared with most other kinds of

North American snakes, the racer is remarkably tolerant of heat, and

often is several degrees warmer than the level that those of other

genera will normally tolerate. Racers have been seen crawling in the

sunshine, or basking on days that were uncomfortably hot for humans.

For example, on August 28, a large female racer released from a trap

was followed and her behavior observed; after crawling some 50 feet

through the grass she climbed from a ditch bank onto sunflower stalks

and elm saplings, and came to rest among the stalks, in a spot

strategically situated for catching grasshoppers. More than half her

body was exposed to sunshine and air temperature was slightly above

34° Centigrade, yet the snake showed no sign of discomfort, and for

the several minutes that she was kept under observation, did not

attempt to withdraw into the shade.

[Illustration: Fig. 6. Bodily temperatures of blue racers captured by

hand in their natural surroundings. The preferred level is

approximately the same as indicated by Fig. 5 (between 29° and 36°

Centigrade), but some of the racers caught were not fully active and

had lower temperatures. Some bias results from the fact that those

having the lowest temperatures were the least active and hence were

most easily caught.]

[Illustration: Fig. 7. Air temperatures recorded at captures of the

racers whose records were used in Fig. 6. An active racer typically

maintains, by basking, a bodily temperature several degrees warmer

than the air.]

[Illustration: Fig. 8. Bodily temperatures of blue racers found in

live-traps at Harvey County Park. Opportunity to regulate temperature

by behavior was limited in these snakes in traps, which tended to

match ambient temperatures.]

At the Harvey County study area, bodily temperatures were recorded in

many of the racers that were caught in traps. These records are much

less significant than the records obtained from racers caught by hand

and promptly checked for bodily temperatures. The temperatures of the

trapped snakes may, to a large extent, reflect the temperatures of air

and soil at the time. However, despite their confinement, the trapped

racers probably were able to exercise some control over their

temperatures by shifting from shade to sunshine, or from the top of

the trap to its bottom, where they would be in contact with the

substrate. In most of the racers removed from traps, as in those

caught by hand, bodily temperatures were somewhat above air

temperatures, but the difference was less in the former group of

snakes. Figure 8 shows the bodily temperatures of these snakes removed

from traps. Occasionally racers died in the traps from overheating. On

July 2, 1960, each of two racers in traps had temperatures of 39.4°.

One of these was especially vicious and frantic in its attempts to

escape, but otherwise seemed unharmed. The second racer was dead,

seemingly having succumbed just before it was found. Probably

prolonged exposure to temperature in excess of 39° would always result

in death of the snake. Racers and other snakes that had become

overheated in the traps and were nearing exhaustion had a

characteristic limp feel when they were handled. In June, 1960, heat

tolerance of a halfgrown racer was compared with that of several other

snakes including a copperhead, garter snakes, and ringneck snakes.

Each snake in turn was enclosed in a plastic tube plugged with cotton

at one end, the snake having a quick-reading thermometer taped in

place for a rectal reading. The tube was then placed in sunshine. Over

periods of minutes the enclosed snake passed through a characteristic

cycle. Soon it would begin to register discomfort as its temperature

rose rapidly. Its struggles would become increasingly violent, then

would cease abruptly. The snake would suddenly collapse, its body

mostly limp, but knotted in slow contortions, its mouth gaping widely.

Within a few seconds all movements would cease, but in each instance

the seemingly dead snake was soon revived by holding it in cold

running water. The copperhead, garter snakes, and ring-necked snakes

all collapsed at temperatures near 41°. At this same temperature the

racer showed signs of acute discomfort, but did not collapse even

after many minutes of exposure. Probably more protracted exposure at

this level would have been fatal to the racer as well as to the other

kinds tested.

Natural History of the Racer Coluber constrictor · 第 33 / 47 题