Thamnophis sirtalis, in North-
f Kansas

Natural History Museum, The University 0

2001. Fitch, H. S. Further study of the Garter Snake,

eastern Kansas. Scientific Papers,

19: 1-6.

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-06369 2 gay Big P AUG 0 6 2001
= 'clentific Papers
| UNIVERSES
Natural History Museum
The University of Kansas
15 June 2001

Number 19:1-6

Further Study of the Garter Snake, Thamnophis sirtalis,
in Northeastern Kansas

By

Henry S. FitcH

Fitch Natural History Reservation, The University of Kagssas
2060 East 1600 Road, Lawrence, Kansas 66044-9450, USA

CONTENTS -

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ABSTRACT The red-sided garter snake, Thamnophis sirtalis parietalis, is a prominent member of the
local snake fauna at the Fitch Natural History Reservation and has been a subject of field study over
the past 53 years. Although several publications in whole or in part, have been devoted to this species,
some aspects of its ecology have remained poorly known; some of these are dealt with in the present
paper. Herein, I discuss the range of variation in color and pattern, and compare the frequencies of
pattern types in males and females; I also compare the sexes with each other and with immatures in
their ecological traits including vagility and their fidelity to specific locations. Comparison of feeding
records show major differences in food habits between the sexes and between young and adults. A

northern population of this same subspecies in the Interlake Region in Manitoba, Canada, is com-
pared behaviorally with the Kansas population on the basis of recent studies.

Key worps: Dorsolateral area; Funnel trap; Intersquamal color; Probability of capture; Thamnophis sirtalis
parietalis.

© Natural History Museum, The University of Kansas ISSN No. 1094-0782

2 SCIENTIFIC PAPERS, NATURAL History MuseuM, THE UNIVERSITY OF KANSAS

INTRODUCTION

The common garter snake, Thamnophis sirtalis, is eco-
logically important and geographically widespread; it oc-
curs throughout most of the United States (except the arid
southwestern one-fourth) and in the southern half of
Canada (Ruthven, 1908). A large amount of literature ex-
ists regarding its local occurrence, identification, and in-
teractions with other species (Fitch, 1980). It is in fact one
of the best known of North American snakes. The recent
book by Rossman et al. (1996) summarizes much informa-
tion concerning the species. My acquaintance with the spe-
cies began with studies of the west coast garter snakes at
the University of California’s Museum of Vertebrate Zool-
ogy in the 1930s.(Fitch, 1941). With my becoming the first
superintendent of the University of Kansas Natural His-
tory Reservation in 1948, I had opportunity to study a lo-
cal population of the red-sided garter snake (Thamnophis
sirtalis parietalis) and subsequently published an ecologi-
cal study on that snake (Fitch, 1965). In that study the popu-
lation on the Reservation was compared with that in
Harvey County State Park, 140 miles to the southwest; geo-
graphic variation over the entire range was discussed, and
the two local populations were compared with respect to
habitat, temperature relationships, food habits, breeding,

growth, movements, defense and escape, natural enemies,
parasites, composition of the population and numbers. In
my synthesis of a Kansas snake community (Fitch, 1999), I
presented data on the common garter snake dealing with
traits of the species, behavior, size relationships, pattern
types, tail functions, spatial relationships, kinds of prey,
reproduction, growth, numbers, and geographic differen-
tiation. Since that study was completed, three more sea-
sons of field work on garter snakes have been finished.
The present report deals with aspects of the ecology not
undertaken or adequately treated in previous publications,
to describe in detail the color pattern phases and their in-
cidence in a large sample, and to compare the sexes with
each other and with immatures in their ecological traits.
ACKBOWLEDGMENTS

I thank William E. Duellman for his advice regarding
the content and organization of this paper. I am indebted
to Neil B. Ford and Stevan J. Arnold for their many help-
ful suggestions. My daughter, Alice F. Echelle, helped me
with various phases of the early field work and also made
suggestions for improving the original manuscript. Also,
I thank the many students who have helped me capture
snakes over the past 53 years.

MATERIALS AND METHODS

Garter snakes were captured in wire funnel traps, or
by hand from beneath shelters, both metal and wooden,
and these contrasting techniques were used on different
areas. Data routinely recorded from each snake captured
included sex, snout-vent length, tail length, color and
markings, and stomach contents. Color pattern was de-
scribed with special attention to the vertebral stripe, the
dark dorsolateral area, and the series of pale intersquamal
marks on the body. Lincoln Index censuses were made from
time to time on relatively small and discrete areas of in-
tensive study. Each census was based on two consecutive
sampling periods; the ratio of marked snakes (from the
first sampling) to unmarked snakes in the second sampling
was the basis for an estimate; it was obvious that the
snakes’ movements prevented a highly accurate census,
and the longer the sampling period the more distorted the
census figure, because previously marked snakes wan-

dered away and were replaced by new ones. Food habits
were studied by palping food items from the stomach up
into the mouth for identification, and comparisons in-
volved male and female snakes, adults versus first-year
young, areas of different habitats and changes over time.

Areas of most intensive study on Fitch Natural His-
tory Reservation (FNHR) included House Field (10 ha in
the headquarters area sampled from shelters in the 1990s),
and Quarry Field (a 3.36 ha hilltop field on the northern
edge of the reservation). On the adjacent Nelson Environ-
mental Study Area (NESA), study areas included the Elec-
tric Mouse Pens (in the northwestern sector, 9.65 ha
sampled with traps in the 1990s), the Fish Pond Pens (2.6
ha south of NESA headquarters, sampled with traps in the
1990s), and Plots 4 and 5 in the southeastern part of the
Biotic Succession Area (5.0 ha , sampled with shelters, 1986-
2001).

RESULTS

Color pattern.—Thamnophis sirtalis has a longitudi-
nally striped pattern, and is one of the most brightly col-
ored snakes. There is much variation, both geographic and
individual, in color and pattern, but ontogenetically there
is little change, which involves a brightening from the
rather dull color of the neonate. Between the scales on loose
skin of the dorsolateral area are the colored marks that give
the subspecies its vernacular name “red-sided.” These

marks are not always red and are highly variable in size
and color, providing a variety of pattern types. Table |
shows the relative frequency of nine pattern types in the
local population and shows differences between the sexes.
Actually, individual variation is such that a continuum
might be established among all nine types. The
intersquamal marks are arranged in transverse alternat-
ing series, with the two lower rows and the three upper

FURTHER STUDY OF THE GARTER SNAKE 3

Table 1. Pattern variation in intersquamous marks of dorsolateral area
in a local population of Thamnophis sirtalis parietalis. Numbers are per-
centages of total sample size for each sex (669 males, 1029 females).

Intersquamous pattern Males Females
All red 26.5 34.5
All yellow 5.5 35)
All green 3.9 Def
All blue 32 1.2
Uppermost row pale (russet,

ivory, green, or blue) 19.0 14.9
Two upper rows pale 26.5 26.6
Three upper rows pale 12.7 9.1
Four upper rows pale 2.5 2.3
Two upper rows missing

(T. s. fitchi pattern) 0.1 0.2

offset and alternating, separated by about the length of
one scale from the next in front or behind. The lowest row
is just above the lateral stripe and its marks are the largest;
those marks farther from the lateral stripe become progres-
sively smaller and less colorful. The two lower marks are
on each side of the fourth scale row; the three upper marks
involve scales of the fifth, sixth, and seventh rows. These
marks on the dorsolateral area are sometimes all one color
(in both sexes combined, 26.3% red, 3.6% yellow, 2.5%
green, 2.2% blue), but in some cases only the lower rows
are red, and the upper (four, three, two or one) are paler,
faintly yellow, green or blue. In occasional individuals,
the upper series (two or three nearest the dorsal stripe)
are missing, and these snakes have essentially the pat-
tern of the West Coast subspecies, Thamnophis sirtalis
fitchi. In the majority of local snakes, the upper marks
are not red but are pale, ivory, or delicately tinted with
yellow, green, or blue. In a small minority, there is no
red in the pattern and all dorsolateral marks are of pale
color.

Table 2. Relative probability of capture of the red-sided garter snake.

The yellow vertebral stripe is perhaps the most char-
acteristic trait of the species, but in the local population it
also is subject to much individual variation. Typically the
stripe is bright yellow and involves the vertebral scale row
and half of the adjacent row on each side, but it may be a
dull, muddy yellow, may be broader than the typical one
and two halves of scale rows, and may have indistinct
edges. The dorsolateral area between the vertebral stripe
and the lateral stripe on each side is typically sepia-colored
or almost black, but may be paler brown or gray, with two
alternating rows of darker spots and five pairs of transverse
markings, largely confined to the skin between the scales.

Movements and abundance.—Behavioral differences
between the sexes affect their relative probability of cap-
ture (Table 2). For example, site fidelity is more prominent
in females. This is illustrated by attempts to apply a Lin-
coln Index census to the data. In 1998 in House Field, snakes
were captured exclusively by hand. There were 43 males
and 55 females (excluding the young born in late summer).
Of the 98 snakes in the sample, 57 were caught in the first
half of the season (March, April, May. June), 41 were caught
in the latter part (July through October) and 10 of these
were captured in both early and late periods. From the
ratio of recaptures in late summer and fall the Lincoln In-
dex indicates a population of approximately 234. But if
the census is applied separately to males and females, the
figures for females are 38 for the first half of the season, 17
for the second half, and 8 for both, with the equation indi-
cating a population of 80.8 individuals. For males, the cor-
responding figures are 19, 24 and 2, suggesting a popula-
tion of 228 individuals. Intuitively, it seems that the figure
for males is much too high, and is the result of only two
recaptures, because most of those present in spring and
early summer had moved away by the time of the late sum-
mer-fall sampling. Censusing from the ratio of marked to
unmarked snakes assumes there is no movement to or from

Number of captures

Method of Twice Three times Four times or more
Area capture Sex Total N %o N % N %o
House Field Under shelters C 175 76 43.4 16 Ox 6 3.4
or by hand Q 184 103 56.0 75 38.7 27 14.7
Biotic Succession Area Under shelters C 46 5 10.9 1 6.5 - -
or by hand ce) 93 24 25.8 11 11.8 ce) ow.
Nelson Environmental Wire funnel (oy 140 35 25.0 15 10.7 4 2.9
Study Area traps Q 140 50 35.7 20 14.3 12 8.6
Quarry Wire funnel fo 35 10 28.6 8 22.9 - -
traps 2 Bil 30 58.8 17 33.3 6 16.8
Hilltop outcrop Wire funnel (oy 53 3} 6.0 = = = =
traps ie) 81 5 6.3 1 1.3 = =
Reservation, Wire funnel toy 457 48 10.5 10 DP 10 2.2
1950s & 1960s traps ie) 635 94 14.8 23 3.6 19 3.0

4 SCIENTIFIC PAPERS, NATURAL History MuseuM, THE UNIVERSITY OF KANSAS

Table 3. Distances between capture points and implied areas of activity
for male, female, and young red-sided garter snakes.

Males Females Young
N 147 398 106
Mean distances (m) between capture points 117.9 116.3 80.6
Implied area (ha) SHY. 3.67 2.23
Recaptures reflecting no net movement 9 44 13

a study area. However, there is always some movement,
depending on the length of time involved. In male garter
snakes movement is especially prominent, and it is essen-
tial to have short sampling periods in order to minimize
the effect (Fitch, 1999) and to be aware of the error caused
by movement. Compared with females, adult males move
farther and more frequently, and they have less tendency
to return to any specific location. Therefore, they are less
susceptible to recapture, and this affects the ratios in Lin-
coln Index censuses. ‘

Table 3 shows differences between the sexes and be-
tween adults and young in distances moved. The young
were all first-year individuals, but most had made sub-
stantial growth before their first capture. They were mostly
in the range of 300-400 mm in snout-vent length, and thus
had about doubled in length since birth. In these partly
grown young, the average movements were substantially
less than in adults of either sex. It is believed that even
smaller young, down to the size of neonates would show
progressively shorter movements and their home ranges
would be progressively smaller. Adults of both sexes are
similar in their movements compared with young, which
move within smaller areas. Adult females and young are
alike (but differ from males) in their tendency to return
frequently to favored spots.

On the FNHR until the 1980s, and at the pens area
in the NESA, samples of snakes were obtained with wire
funnel traps and consisted almost entirely of adults,
because immatures easily pass through the quarter-inch
wire mesh of the funnel traps. In the Biotic Succession
Area (BSA) of the NESA and FNHR in the late 1980s
and 1990s, sampling was accomplished by distributing
wooden and metal shelters, which presumably were
used by snakes of all sizes and ages indiscriminately,
but young were relatively few in the upland habitat of
the BSA.

Food habits.—Studies of food habits have shown
major geographic differences within the species but usu-
ally have not compared food of the sexes, or compared
adults and young. In the local population such differ-
ences exist and are shown in Tables 4 and 5. Table 4
shows the total food sample divided into 48 categories
to bring out differences between the sexes, between
adults and young, and between areas of different habi-
tat.

There are striking differences between areas. The
many young captured in House Field had eaten earth-
worms almost exclusively. One had eaten a small slug
(Deroceras Iaeve) and several had eaten newly metamor-
phosed hylids (Acris and Pseudacris). Another impor-
tant difference was predation on small mammals by
adult females (30 records, 16 of them on Muicrotus
ochrogaster) compared with only one record for a male
snake, which had eaten an individual of the much less
bulky Reithrodontomys megalotis.

Frogs (Rana blairi and R. catesbeiana) made up a major
part of the food and were well represented in each local-
ized sample. Rana blairi was eaten about twice as often as
R. catesbeiana, and none of the latter species was much more

Table 4. Prey of the red-sided garter snake in four areas of the Fitch Natural History Reservation (FNHR) and Nelson Environmental Study Area
(NESA) comparing sexes, mean snout-vent lengths inmm (SVL + 1 SE), and numbers (N) of snake predators. BSA = Biotic Succession Area of NESA.

House Field FNHR BSA NESA Pens
1980-2000 1950-1990 1986-2000 1986-2000
Prey Sex N SVL N SVL N SVL N SVL
Earthworm (oy 54 397.9 +121 9 517.7 2 356 - =
ie) 81 430.6 + 11.3 4 475.8 + 21.1 6 480.8 + 16.9 517
Microtus C = = = = = = == =
2 1 623 3 675 6 649.3 + 32.7 6 705.7 + 21.5
Other mammals* (oy = = 1 490, - = - =
ie) 4 600.8 + 28.8 10 636.4 + 3.14 1 783 4 612.3+8.8
Rana* oy 1. 496.3 41.5 15 488.9+2.5 1 540 6 536.7 + 6.2
ie) 23 567.6+4.3 21 549.0 + 6.7 4 669.3 + 14.4 27 623.0 + 13.8
Bufo* (oy 3 438.6 6 403..3 + 0.65 - ~ 2 374.0
ie) 3 582.4 7 615.3 + 6.2 4 673.0 + 18.5 1 745
Other amphibians* Cy 3 354.5 4 364.8 + 14.1 4 442.5 +74.2 3 338.3
Q 9 442.2 7 562.4 + 10.0 5 532.6 +12.1 9 591.4+3.9

**“ Other mammals” includes Peromyscus, Reithrodontomys, Blarina, and Cryptotis.

“Rana” includes R. blairi and R. catesbeiana. “Bufo” includes B.

americanus and B. woodhousti. “Other amphibians” includes Acris, Hyla, Pseudacris, and Gastrophryne.

FURTHER STUDY OF THE GARTER SNAKE 5

Table 5. Prey eaten by red-sided garter snakes on the FNHR and adjoin-
ing areas and sizes of snake predators. In columns of SVL, the first line
is the observed range and the second line is the mean + 1 SE. Measure-
ments in mm.

SVL of male SVL of female

Kind of prey* N predator N predator

Allolobophora caliginosa 60 225-666 88 226-666
(earthworm) SoS) se 17 432.1+10.5

Rana blairi 47 411-610 54 363-780
Sp} a2 C)5) 602.9 + 13.3

Rana catesbeiana 7 419-582 13 531-834
487.4 + 23.4 637.8 + 32.7

Rana sp. 1 497 8 294-630
= 554.9 + 46.8

Bufo americanus 5 368-596 13 519-745
458.0 + 52.8 638.5 + 52.9

Bufo woodhousii 5 459-578 6 488-673
432.6 +40.1 616.7 + 40.4

Hyla chrysoscelis 5 243-596 9 374-653
426.6 + 63.9 571.7 £ 30.5

Acris crepitans 7 228-580 10 240-712
413.7 55.2 480.6 + 59.8

Pseudacris triseriata 2 383-406 4 303-688
394.0 487.3 + 96.0

Microtus ochrogaster = = Y/ 585-810
= 691.1 + 23.6

Peromyscus leucopus - - 8 510-850
= 647.3 + 44.1

Reithrodontomys megalotis = = 5 565-783
- 637.4 + 37.9

*Prey species with few records (sex and SVL of snake predator in paren-
theses): Cryptotis parva (Q, 615), Synaptomys cooperi (9, 648), Cardinalis
cardinalis (Q, 713), Melospiza lincolnii (Q, 520). Unidentified bird (@, 634),
Diadophis punctatus (, 520), Gastrophryne olivacea ( 9, 600),Deroceras laeve
(2, 303).

than half grown. Toads (Bufo) were taken about half as of-
ten as frogs (Rana), and several of those eaten, especially
of the smaller B. americanus, were adults. Of the tree frogs
(Hyla chrysoscelis) eaten, several had been taken by snakes
found at the hilltop rock outcrops where the snakes hiber-
nate. Perhaps the frogs also had come to these places in
search of hibernacula.

Ina total of 375 prey animals, most were amphibians
(8 kinds), small mammals (6 kinds) and earthworms, with
an occasional bird, snake, or slug. Food of the first-year
young consisted almost exclusively of earthworms, but
with an occasional newly metamorphosed anuran. Of 32
small mammals eaten, 31 including all the microtines (13)
were taken by adult females (510 to 850 mm SVL) and only
one mammal, a harvest mouse (Reithrodontomys) was eaten
by an adult male (490 mm SVL). Frogs (Rana) and toads
(Bufo) were eaten chiefly by the adult snakes, especially
by females. It should be emphasized that mammals, al-
though too bulky to be ingested by males and immatures,
are regularly eaten by adult females and constitute an im-
portant resource for the developing embryos.

DISCUSSION

The widespread occurrence of Thamnophis sirtalis
parietalis is evidence of its plasticity and ability to thrive
under diverse conditions. In recent decades, the snake dens
of Manitoba Canada, have attracted much attention. There,
field studies of far northern representatives of T. s. parietalis
by Aleksiuk (1976), Aleksiuk and Stewart (1971), Crews et
al. (1988), Gregory (1974), Joy and Crews (1988), Shine et
al. (2001), and Shine and Mason (2001) have revealed the
following traits:

1. Formation of huge denning aggregations (Joy and
Crews, 1985; Crews et al., 1984); thousands of snakes hi-
bernate together in deep limestone sinks.

2. Formation of massive mating aggregations at, or
near, the hibernacula, as recently emergent males pile up,
sometimes by the hundreds, to court emerging females
(Gregory, 1977).

3. Presence of “she-males” in mating aggregations
(Shine and Mason, 2001); these are the most recently
emerged males, weak and debilitated from their long so-
journ underground, but possessing the ability to produce
pheromones like those of emergent females that are highly
attractive to males (Shine et al., 2001). Within a time-span
as short as one day such males lose much of their attrac-
tiveness but gain in strength and vigor.

4. Massive mortality at and near the dens (Shine et al.,
2001). Predators, including crows, kill many, but much
larger numbers are victims of their own social system. The
pile ups that occur in mating aggregations may suffocate
hundreds of the participants. Mortality is selective; males
that are eliminated tend to be the smallest, thinnest, and
weakest of those participating, and their deaths may ben-
efit the population at large. However, the females killed
tend to be the largest and heaviest, as these are the most
attractive and summon the largest retinues of males.

5) Long distance dispersal (up to 20 km) from the dens,
as the emergent snakes scatter to exploit extensive marsh-
lands in the region of the dens, where they feed on a vari-
ety of marshland animals, notably the wood frog, Rana
sylvatica (Gregory and Stewart, 1975).

It must be realized that these combined traits are not
characteristic of the subspecies T. s. parietalis but, instead,
result from the unique selective pressures generated by
the environment near the northern edge of the range. None
of the behaviors described above has been observed in the
red-sided garter snakes of northeastern Kansas, thereby
indicating widely divergent responses to contrasting en-
vironments. No comparison of morphology or color pat-
tern has yet been made between the Kansas population of
T. s. parietalis and that of Manitoba.

The longitudinally striped pattern is adaptive in that
it conceals motion as the snake moves through dense veg-
etation. It is significant that adult females, larger and more

6 SCIENTIFIC PAPERS, NATURAL History MusEuM, THE UNIVERSITY OF KANSAS

aggressive than male counterparts, are somewhat more
likely to have red in the pattern and to respond to con-
frontation with sematic display, whereas males are less
aggressive and more inclined to escape and to seek con-
cealment.

Shine (1991) pointed out that Thamnophis are unusual

among snakes for their marked sexual dimorphism in head
size, with the females having relatively larger heads. Mam-
mals, although too bulky to be ingested by males and
immatures, are eaten regularly by adult females. This di-
etary difference helps to explain how this sexual dimor-
phism is maintained.

LITERATURE CITED

Aleksiuk, M. 1976. Reptilian hibernation. Evidence of adaptive strate-
gies in Thamnophis sirtalis parietalis. Copeia 1976:170-178.

Aleksiuk, M., and K. Stewart. 1971. Seasonal change in the body compo-
sition of the garter snake (Thamnophis sirtalis parietalis) at northern
latititudes. Ecology 52:485-490.

Crews, D., M. Diamond, R. Tokarz, B. Camazine, and W. Garstka. 1984.
Hormone independence of male sexual behavior in a garter snake.
Hormonal Behavior 18:29-41.

Crews, D., V. Hingorani, and R. J. Nelson. 1988. Role of the pineal gland
in the control of annual reproductive, behavioral, and physiologi-
cal cycles in the redsided garter snake (Thamnophis sirtalis parietalis).
Journal of Biological Rhythms 3:293-302.

Fitch, H.S. 1941. Geographic variation in the garter snakes of the species
Thamnophis sirtalis in the Pacific Coast Region of North America.
American Midland Naturalist 26:570-592.

Fitch, H. S. 1965. An ecological study of the garter snake, Thamnophis
sirtalis. University of Kansas Museum of Natural History 15:493-564.

Fitch, H. S. 1980.Thamnophis sirtalis. Catalog of American Amphibians
and Reptiles 279:1-4.

Fitch, H. S. 1999. A Kansas Snake Community: Composition and Changes
over Fifty Years. Malabar, Florida: Krieger Publishing Company, xi
+ 165 pp.

Gregory, P. T. 1974. Patterns of spring emergence of the red-sided garter
snake (Thamnophis sirtalis parietalis) in the Interlake region of
Manitoba. Canadian Journal of Zoology 52:1063-1069.

Gregory, P. T. 1977. Life history parameters of the red-sided garter snake
(Thamnophis sirtalis parietalis) in an extreme environment, the
Interlake region of Manitoba. National Museum of Canada, Publi-
cations in Zoology 13:1-44.

Gregory, P. T., and K. W. Stewart 1975. Long-distance dispersal and feed-
ing strategy of the red-sided garter snake (Thamnophis sirtalis
parietalis) in the Interlake of Manitoba. Canadian Journal of Zool-
ogy 53:238-245.

Joy, J. E., and D. Crews. 1985. Social dynamics of group courtship behav-
ior in male red-sided garter snakes (Thamnophis sirtalis parietalis).
Journal of Comparative Psychology 99:145-149.

Rossman, D. A., N. B. Ford, and R. A. Seigel. 1996. The garter snakes:
evolution and ecology. Norman, Oklahoma: University of Oklahoma
Press, xx + 331 pp.

Ruthven, A. G. 1908. Variation and genetic relationships of the garter-
snakes. Bulletin of the United States National Museum 61, xii +
201 pp.

Shine, R. 1991. Why do larger snakes eat larger prey items?.Functional
Ecology, 5:491-502.

Shine, R., M. P. LeMaster, L. T. Moore, M. M. Olsson, and R. T. Mason.
2001. Bumpus in the snake den: effects of sex, size and body condi-
tion on mortality of red-sided garter snakes. Evolution 55:598-604.

Shine, R., and R. Mason. 2001. Serpentine cross-dressers. Natural His-
tory Magazine. February 2001:59-61.

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ral History Museum, The University of Kansas, Lawrence, Kansas 66045-2454, USA.
Purchasing information can be obtained by calling (785) 864-4450, fax (785) 864-
5335, or e-mail (kunhm@ukans.edu). VISA and MasterCard accepted; include ex-
piration date.

Series Epiror: William E. Duellman

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