Summary

Field study of the blue racer was carried on in several localities in

Kansas, but chiefly at the University of Kansas Natural History

Reservation (the northeasternmost section of land in Douglas County),

and the adjacent 160-acre Rockefeller Experimental Tract in Jefferson

County. By October 26, 1962, after 14 years of field work, a total of

1423 racers had been captured some 2197 times.

The locale of the present study was near the geographic center of the

blue racer's range. The range, chiefly in the Mississippi Valley and

Great Plains region, is centrally situated with respect to the other

seven subspecies. An extensive but scattered literature concerning the

ecology of the species as a whole, and its several geographic races,

has been reviewed and utilized for comparison with my own field data.

Blue racers were caught in wire funnel traps set in prairie and

pastureland habitat in summer, and along hilltop limestone outcrops in

woodland in autumn. The autumn trapping along rock outcrops was

carried on each year from 1949 to 1962, but effective summer trapping

was carried on only in the last six years of the study. Each racer

caught was individually and permanently marked by scale clipping. More

than half were caught only once, but many were recorded repeatedly,

with a maximum of 16 captures.

The racer occurs throughout most of the United States, and its

populations are subject to much geographic variation. The snakes are

largest in the northeastern part of the range, with clines of

decreasing size toward the southeastern, southern and western parts of

the United States. There are somewhat parallel trends in coloration;

the black racer of the northeastern states grades into paler, gray or

light brown subspecies in southern Florida, Texas, and the far western

states. Accompanying these changes in color and size are minor

morphological changes and major ecological changes. The black racers

of the eastern states often inhabit forest or forest-edge habitats

while the paler and smaller snakes of more southern or western areas

typically inhabit scrub, chaparral, or prairie. The large,

dark-colored racers of the eastern and northeastern states are

especially inclined to attack larger prey including small

vertebrates, even weasels, rabbits, and chipmunks, whereas the

smaller and paler racers of more southern and more western areas take

a higher proportion of insects and rarely attack vertebrates other

than small reptiles.

On the area where field work was carried on in northeastern Kansas,

tall-grass prairie habitat is preferred, but fields of grain or

alfalfa, grazed pasture, brush, woodland edge, groves or open

woodland, and weedy fields are all utilized to some extent. The racer

is strictly diurnal and largely terrestrial but it may climb through

bushes or small trees in foraging or escaping.

The blue racer is a typical colubrine snake of slender build, with

large eyes, and vision plays an important role in finding prey and

detecting enemies. In the adult blue racer the dorsal color is

variable, pale brown or gray, bluish, greenish or slaty. In the

hatchling, however, there is a distinct pattern of a type widespread

among colubrines and also among snakes of other groups--a series of

middorsal blotches on an olive ground color, with alternating rows of

smaller spots on each side. The ventral surface is pale, with dark

speckling. The pattern is sharply defined on the anterior part of the

body, but markings become progressively more obscure posteriorly and

are scarcely discernible on the tail.

The juvenal pattern fades gradually as growth proceeds, and there is

much individual variation in the rate of its loss. Some racers still

retain the juvenal pattern faintly discernible after attainment of

sexual maturity. There are also striking ontogenetic changes in the

proportions of the head, body and tail. The diameter of the eye is

approximately one per cent of the snout-vent length in hatchlings, but

is only a little more than half that relative size in the largest

adults. In the course of allometric growth other parts of the head

also enlarge less rapidly than the body, but more rapidly than the

eye. In hatchlings there is a slight average difference between the

sexes in relative tail length, with males' tails the longer. Relative

tail length increases slightly in both sexes up to the time of sexual

maturity, and then decreases slightly with advancing age.

Racers in northeastern Kansas spend nearly half the year in

hibernation, with average recorded emergence date April 16, and

average date of retirement into hibernation November 8. Hibernacula

are usually in crevices in hilltop limestone outcrops with south

exposures. Winter temperatures within the hibernacula are usually well

within the range 0 degrees to 10 degrees Centigrade. Spring emergence

has been recorded at an air temperature of only 12.5 degrees

Centigrade. Racers bask in sunshine frequently even in warm weather,

and the temperature preferendum is several degrees higher than in most

other kinds of snakes. Bodily temperatures obtained from blue racers

that were fully active, either under natural conditions or in a large

outdoor enclosure, were concentrated in the neighborhood of 34 and 35

degrees Centigrade. For short periods racers can survive temperatures

up to 45 degrees without damage, but more prolonged exposure to

temperatures of slightly less than 40 degrees can be fatal. In

hibernation, racers can withstand temperatures slightly below

freezing, but they cannot survive being frozen solid.

Blue racers tend to limit their activities to familiar areas or home

ranges; some individuals may live out their entire lives within the

same home range, but others shift from time to time. Average home

ranges of approximately 26 acres for males and 24 acres for females

were calculated. The racers' preference for hibernacula in a habitat

different from that to which summer activities are confined

necessitates spring and fall migrations between the limestone outcrops

where hibernation occurs and the grasslands where the snakes stay in

summer. The average spring or fall migration is approximately a

quarter of a mile, but an individual racer does not consistently

return to the same hibernaculum. Many racers were recorded to have

made movements of 2000 to nearly 4000 feet, involving shifts in home

range, but some later shifted back to their original areas. Some may

have made even longer shifts but their movements would not have been

recorded since they would have gotten beyond the limits of the study

area.

Blue racers hunt by various methods, often by coursing through dense

vegetation in active search in which vision is of primary importance

in locating the prey. Almost any small animal that moves nearby may be

overtaken and caught with a sudden dash. From analysis of scats and

prey from stomachs, a total of 1357 food items of more than 50 species

was compiled. Favorite prey species were the cricket (_Gryllus

assimilis_), grasshoppers (_Arphia simplex_, _Melanoplus

femur-rubrum_, _M. bivittatus_, _M. differentialis_), camel crickets

(_Ceuthophilus_ sp.), katydid (_Neoconocephalus robustus_), vole

(_Microtus ochrogaster_), white-footed mouse (_Peromyscus_ sp.),

racerunner, (_Cnemidophorus sexlineatus_), and leopard frog (_Rana

pipiens_). The insects taken greatly outnumbered the vertebrates, but

the vertebrates made up most of the actual bulk of prey eaten.

Crickets, grasshoppers, and katydids comprised most of the insect

prey. Beetles, moths and cicadas were rarely taken. Vertebrate prey

included miscellaneous small snakes (some of them juvenal racers),

mammals, and birds. Seasonal change in the composition of the food is

slight, but vertebrates figure more prominently in early summer, and

insects comprise increasing percentages later in the season.

Composition of the food differs according to size of the snake;

gryllid and ceuthophilid crickets are best represented in the food of

juveniles whereas small mammals, and grasshoppers of the genus

_Melanoplus_ are best represented in the food of large adults.

The breeding season is mainly in May. Mating is promiscuous and two or

more males may court the same female simultaneously. A courting male

lies on or alongside a receptive female, with spasmodic rippling

abdominal movements, and with his vent adpressed to hers. At intervals

in the courtship period the female moves swiftly for a few feet or a

few yards shifting to a new spot, and during her activity the male

strives to maintain contact with her. From time to time the male

leaves the female briefly and courses rapidly around her in a devious

route. Courtship is consummated when the female raises her tail in

acceptance of the male and intromission is effected. During coitus,

which lasts for periods of minutes, the female moves forward slowly,

dragging the passive male, tail-first behind her.

Ovulation normally is in late May. Eggs from 29 to 39 millimeters in

length and 14 to 21 millimeters in breadth are laid, from mid-June to

early August, usually in tunnels of fossorial mammals such as voles or

moles, at depths of five to 12 inches. Clutches of the blue racer

average 11.8 eggs but the number is correlated with age and size of

females; two-year-olds average only 9.2 eggs, whereas those females

that are six years old or more average 15.7 eggs. Also, there is

geographic variation in size of clutch, from only 5.8 eggs in _C. c.

mormon_ of the West Coast to 16.8 eggs in _C. c. constrictor_ of the

northeastern states. In each breeding season some females of adult

size do not produce clutches. Only about 13 per cent of the

two-year-olds in a small sample were fecund, but the ratio increased

to 80 per cent in old adults. Incubation averages 51 (43 to 63) days.

Hatchlings usually make several longitudinal slits in the eggshell

with the egg tooth before emerging, and often require a day or more to

emerge after the first slit is made. Hatchlings average 10-3/4 inches

and 4.16 grams. By late October when these young are ready to enter

hibernation, they have grown to 16-3/4 inches and 12.3 grams. Typical

October lengths (overall) in inches for males and females,

respectively, after successive seasons of growth, are as follows:

yearlings, 27-3/4 and 29; two-year-olds, 34-1/4 and 37-1/4;

three-year-olds, 37-3/4 and 41-3/4; four-year-olds, 40 and 44-3/4;

five-year-olds, 41-1/2 and 46-1/4; six-year-olds, 42-1/4 and 48-1/4;

seven-year-olds, 43-3/4 and 50; eight-year-olds, 44 and 51-1/4.

Judging from trends in the small samples available, sex ratio in

hatchlings is approximately 1:1. In the summer trapping of blue

racers, males are caught in larger numbers than females, but seemingly

this is because of their greater activity. In fall along the limestone

outcrops where the racers hibernate females were caught in slightly

greater numbers than males among the young adults, and made up a

little more than 60 per cent of the old adults. Presumably the males

are eliminated more rapidly, because of their greater activity, their

smaller size, or a combination of both factors. The composition by age

groups of the adult racers captured was as follows: two-year-olds,

41.5 per cent; three-year-olds, 17.8 per cent; four-year-olds, 12.6

per cent; five-year-olds, 9.5 per cent; six-year-olds, 6.1 per cent;

seven-year-olds, 4.3 per cent; eight-year-olds, 2.7 per cent;

nine-year-olds, 2.4 per cent; ten-year-olds, 1.2 per cent; more than

ten years old, 1.9 per cent.

Attempts to census blue racers on the study plots, from the

capture-recapture ratios, yielded highly variable figures for

different areas and even for the same area at different times.

Variability is believed to result mostly from the small sizes of the

samples, none of which is large enough to yield a census figure that

is statistically reliable. Combined samples yielded figures indicating

populations of one to three adults per acre in early summer in areas

of favorable habitat. To represent the entire population in late

summer--the time of its annual maximum--these figures would need to be

approximately tripled.

The blue racer depends mostly on speed to escape its enemies. In

escaping it often takes advantage of a downhill slope to increase its

speed. A racer startled at close range often sets out with a violent

thrashing that attracts attention to a given spot, then glides away so

silently and rapidly that its course is not detected. The snake may

circle back and approach the scene of disturbance slowly and

stealthily, from the direction opposite to that taken in departing.

Often a racer seeks concealment by climbing into a bush or tree, and

occasionally escape is effected by swimming. A racer that is caught or

cornered usually makes a spirited defense by striking and biting. An

alternative reaction, seen most typically when the snake is

handicapped by injury or by low temperature, is to coil with the head

concealed, and, with writhing movements, to smear the surface of the

body with musk discharged from glands in the base of the tail.

Many natural enemies prey upon the racer, but in the present study the

majority of records pertained to the red-tailed hawk. The broad-winged

hawk, marsh hawk, red-shouldered hawk, sparrow hawk, and barn owl also

are among the raptors that feed on this kind of snake. Among mammals

the striped skunk is the only species definitely recorded to prey on

the racer, feeding on the eggs as well as upon the young and adults.

The common king snake, prairie king snake, timber rattlesnake,

copperhead, and even the slender glass lizard have been recorded to

prey on the blue racer, but probably all are of minor importance as

natural enemies. Shrews (_Blarina_, _Cryptotis_) and mice

(_Peromyscus_, _Reithrodontomys_) have sometimes killed and eaten

racers confined in traps, and under natural conditions they possibly

prey upon snakes that are immobilized when torpid from cold, in their

hibernacula and temporary shelters. Chiggers are the most common

ectoparasites of the racers. Four species of the chigger genus

_Trombicula_ have been recorded on racers from the Reservation. The

racer is an important host of the common pest chigger, _Trombicula

alfreddugèsi_, which often attacks humans. Most adult racers on the

Reservation and nearby areas carry the fluke, _Neorenifer

lateriporus_. In spring these flukes are conspicuous in the mouths of

the racers, but in late summer they are not in evidence.

Natural History of the Racer Coluber constrictor · 第 42 / 47 题