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Straightness-of-path during and after vernal migration in prairie rattlesnakes,

Crotalus viridis, in eastern Colorado

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Herpetology Notes, volume 7: 425-436 (2014) (published online on 5 July 2014)

Straightness-of-path during and after vernal migration

in prairie rattlesnakes, Crotalus viridis,
in eastern Colorado

David Chiszar1, Bryon K. Shipley2,*, Hobart M. Smith3, Kevin Fitzgerald4

and Anthony J. Saviola5

Abstract. Nineteen free-ranging prairie rattlesnakes, Crotalus viridis, (ten males, nine females) were tracked with radio
telemetry at the Plains Conservation Center in Arapahoe County, Colorado in 2005 and 2006. The average angle of all
movement segments showed that straightness-of-path was greatest during the vernal migration and declined significantly
afterwards. However, we found no scattered random movements during late spring or summer, indicating that snakes moved
in “directed” ways. Direction might be influenced by external cues, such as landmarks and chemicals or it might be based on
memory or other interoceptive mechanisms. Furthermore, longer movements had higher r-values than shorter ones. Although
only five snakes were followed through the autumnal migration, these showed a return to higher r-values comparable to those
seen during the vernal migration. Hypotheses regarding causal mechanisms underlying rattlesnake movement are discussed
along with proposed experimental tests. Although our r-values were significantly above zero, they were never as high as those
reported for Wyoming conspecifics migrating from hibernacula in the Haystack Mountains, nor were our migrations as long as
those seen in the Wyoming snakes. These differences are attributed to differences in prey distributions in the two ecosystems,
a distinction with conservation implications, especially when captive-raised, head-started, or otherwise provisioned individuals
are to be released into natural habitats.

Keywords. Crotalus viridis, Cynomys ludovicianus, migration, movement, telemetry, straightness-of-path.

Introduction seasonal patterns of migration, as well as movements

related to prey locations, mate searches, and
Understanding movement patterns in regards to
reproductive behaviors. Measurements of home ranges,
habitat and resource availability can provide significant
travel distances and directions, and straightness-of-
insight on the utilization of an animal’s habitat. Further,
path provide useful ecological information about snake
movements in response to foraging or environmental
movement (Gregory, Macartney and Larsen, 1987).
constraints may drastically increase risk to predation as
However, varying habitats and resource availability
well as prove to be energetically costly for migrating
may lead to distinct movement patterns of conspecifics
individuals. Detailed radio telemetry studies in
in different geographical locations. Movements of
rattlesnake ecology have illuminated differences in
prairie rattlesnakes (Crotalus viridis; nomenclature
follows Crother, 2008) in the United States have been
especially well studied (see Jorgensen, Gates and
Whiteside, 2008), although to a much lesser degree
throughout the Great Plains region. Shipley et al.
1
Department of Psychology, University of Colorado, Boulder, (2013) investigated movements of an eastern Colorado
CO 80309, USA population of C. viridis in conjunction with Black-
2
Tropical Discovery, Denver Zoo, Denver, CO 80205, USA. tailed prairie dog (Cynomys ludovicianus) colonies
3
Department of E. E. Biology, University of Colorado, Boulder,
revealing movements that were short (mean = 648 m),
CO 80309, USA
relatively straight, with frequent changes in direction.
4
Alameda East Veterinary Hospital, Denver, CO 80427, USA
5
School of Biological Sciences, University of Northern However, Duvall, King and Gutzwiller (1985) showed
Colorado, Greeley, CO 80639, USA that C. viridis in Wyoming exhibited a remarkable
*
Corresponding author; e-mail: crvi@comcast.net straightness-of-path while moving significantly farther
426
distances (mean = 5,640 m) from hibernacula to their
hunting grounds (i.e., during the spring migration),
likely resulting in considerable energy economy during
the spring migration and probably in other benefits as
well (Brown, 1990; King and Duvall, 1990).
The present paper provides a more in-depth
examination of straightness-of-path in migratory
movements of a population of C. viridis from Colorado
already studied by Shipley et al. (2013). Specifically,
we assessed two factors that have not been studied
previously in any snake: (1) the degree to which the
paths depart from more-or-less straight lines after the
spring migration, and (2) the rate at which the change
occurs. It seems likely that these behaviors ought to be
influenced by the density and distribution of rodents or
other prey. Although rodent surveys were not conducted
in the current study, our results might represent a
baseline situation in that prey in our study area may
be more abundant due to C. ludovicianus activities
(Shipley et al., 2013). Therefore, snake movements in
habitats with lower prey abundance could be longer
and exhibit greater straightness-of-path than we see in
eastern Colorado (Duvall, King and Gutzwiller, 1985).
The only other study of C. viridis movements in the
Great Plains region (Ludlow, 1981) in the presence
of C. ludovicianus was observational, and employed
neither radio telemetry nor capture-mark-recapture
techniques, and therefore no inferential statistics were
reported for traveling distances or straightness-of-path.
The present population of C. viridis lives in association
with C. ludovicianus. Ecological relationships between
C. ludovicianus and reptiles and amphibians have
been studied previously (Reynolds, 1979; Clark et al.,
1982; Kretzer and Cully, 2001; Lomolino and Smith,
2003; Shipley and Reading, 2006). However, snake
movement was rarely the primary objective of such
work (but see Holycross, 1993), yet these papers reveal
that prairie dog colonies provide several important
resources for C. viridis, as is also true in our study area.
For example, in many western locations, including the
present study site, C. ludovicianus burrows provide
hibernacula for rattlesnakes as well as for other reptile
species. Neonates of C. ludovicianus are sometimes
taken by C. viridis, but of greater importance is the fact
that diversity of other small mammals is increased in the
vicinity of active C. ludovicianus colonies (Reading et
al., 1989; Ceballos, Pacheco and List, 1999; Tyler and
Shackford, 2002; Smith and Lomolino, 2004; Shipley
and Reading, 2006). These are the principle reasons for
expecting C. viridis migratory movements to be shorter
David Chiszar et al.
at our site than at the Haystack Mountains of Wyoming
where montane rock outcrops provide hibernacula far
from hunting grounds, where prey are relatively scarce
and widely dispersed in the prairie below (Duvall, King
and Gutzwiller, 1985).
Materials and Methods
We conducted our study at the Plains Conservation
Center (PCC) in Arapahoe County, Colorado (-
104.7364 N, 39.6556 W; datum: WGS84; Figure 1),
although some rattlesnakes occasionally moved beyond
the legal boundaries of PCC. The site is composed
of 445 ha of clayey, loamy, short-grass steppe, with
elevations ranging from 1,728­-1,783 m (Shipley et al.
2013). There are no rock outcrops or arroyos (Figure 2).
This factor forces C. viridis within PCC to use rodent
burrows, especially those of C. ludovicianus, as refugia.
Three C. ludovicianus colonies exist within PCC (7.8,
10.4, and 25.9 ha, respectively) and the colonies are
separated from each other by 1.3-2.4 km.
Adult C. viridis were captured at C. ludovicianus
burrows within all three colonies in April 2005 (eight
males, five females) and 2006 (two males, four females).
Mean mass for males was 520 + 50.9 g (mean + SE) and
for females was 486.7 + 53.1 g. Mean SVL (snout to
vent length) for males was 87.7 + 2.8 cm and for females
was 83.9 + 3.3. Animals were transported to Alameda
East Veterinary Hospital, injected with a passive
integrated transponder chip (HomeAgain™ Microchip
identification system, Schering-Plough Animal Health
Corporation, Omaha, Nebraska, USA), and received an
implant of a G3-1V transmitter (AVM Instrument Co.,
LTD., Colfax, California, USA). Transmitters were
placed subcutaneously in the left side, approximately
midway along the SVL following Fitzgerald and Vera
(2006). The entire package never exceeded 4.5% of
a snake’s mass. Within 24 hours following surgery,
snakes were released into the burrow from which they
were captured. Of the nine females tracked in 2005 and
2006, five were gravid (G) and four were non-gravid
(NG).
Radio tracking began the day after snakes were
released, using a telemetry receiver (LA 12-Q) and a
Yagi 3-element antenna (AVM Instrument Co., LTD.,
Colfax, California, USA). Locations were recorded as
GPS coordinates using a hand-held receiver (Garmin
eTrex Venture Cx, Garmin International Inc., Olathe,
Kansas, USA). Snakes were located 3-5 times per week
throughout the active season or until the signal was lost,
usually through battery failure but sometimes because
Straightness-of-path during and after vernal migration in Crotalus viridis
Figure 1. Map of the Plains Conservation Center (PCC) in Arapahoe County, Colorado, USA.
Figure 2. Landscape view of the Plains Conservation Center
(PCC) in Arapahoe County, Colorado, USA.
427
of predation on the snakes. Five snakes were followed
through the autumnal migration back to hibernacula.
Snakes were located an average of 32 times, so that
detailed tracks could be plotted for each snake on
maps of the study area. Furthermore, the successive
location data were analyzed by circular statistical
methods (Batschelet, 1981; Fisher, 1993) to quantify
straightness-of-path. The principle descriptive statistic
is r (measure of angular dispersion), which takes values
between 0 and 1. When the r-value = 1, all locations
of a given individual lie exactly on the same straight
line, as they all have the same bearing relative to due
north (N). However, when the r-value = 0, locations
may be scattered in a random manner (e.g., a “random
walk” pattern), or they may be balanced such that some
number of locations deviating in one direction from N
are balanced by a similar number of locations in the
opposite direction (e.g., a zigzag pattern of movement
428
in which vectors at one angle to N are balanced by the
same number of vectors at the opposite angle). Either
case represents an absence of concentration along a
constant bearing. Since we have never seen systematic
zigzag movement in C. viridis, we interpret near zero r-
values as random changes of direction, as when a snake
might adopt several ambushing stations sequentially,
each facing a traditional rodent trail from a different
direction.
We defined the spring migration as complete when a
snake had two consecutive days with movements of less
than 10 m. These two days constitute the start of the
post-migration data set for each snake. One male was
lost on June 2nd 2006, four males and two females were
tracked through all or most of July, three males and four
females were tracked through all or most of August, and
two males and three females were tracked until entering
hibernation in late September or October.
Data were analyzed with R version 2.15.2 using
Analysis of Variance (ANOVA) to examine duration,
daily distance movement, and straightness of path (r-
values) between males, NG females, and G females.
Paired t-tests were applied to r-values following the
migration for all three groups of snakes to examine if
there was a significant reduction in straightness of path
after migration. Prior to parametric tests, distributional
properties (i.e., frequencies) of observed scores were
compared with expected frequencies under the null
hypothesis of normality. In addition, non-parametric
(Chi square, χ2) tests were applied to compare male and
pooled (gravid and non-gravid) female data sets where
applicable. Goodness-of-Fit tests were used to make
these comparisons; in no case was the null hypothesis
rejected.
In addition, it is possible that the value of ordinary
means of r or any other index of behavior plotted over
successive days, weeks, or months becomes increasingly
suspect as fewer and fewer subjects contribute data.
Therefore, Vincent (1912) developed a statistical
technique that addressed this problem (see Jiang,
Rouder and Speckman, 2004; Rouder and Speckman,
2004). Each subject’s data record was divided into
successive equal-size fractions, say successive fifths. In
the present case, imagine a data record of 30 successive
location coordinates, such that each successive fifth
would contain six scores. Each segment of six scores
is converted to an r-value, so that the subject ends up
with five successive r’s. This is done for each snake,
dividing its total number of scores into five segments
and taking the r-value for each segment, giving us an
David Chiszar et al.
equal number of Vincentized scores for all animals.
Thus, we can present a summary of mean r-values
across Vincentized blocks. This method does not solve
all problems associated with data loss over time, but
at least it provides a summary in which all subjects
contribute data to each successive block. An important
requirement for the calculation of r-values is that no
less than three location coordinates must exist in each
snake’s five data segments (Batschelet, 1981; Fisher,
1993); in the present study no snake had less than four
locations per data segment.
Vincentizing the movement and r-values during late
spring and early summer allows us to identify which
subject contributed to each successive block. That, in
turn, permitted various statistical analyses that could not
otherwise be applied because of data and subject loss
over the active season. However, the Vincent procedure
has some limitations (Jiang, Rouder and Speckman,
2004; Rouder and Speckman, 2004). The most
important is the fact that the procedure can force snakes
in different motivational states into the same Vincent
block. Consider a male whose signal was lost early in
the season before he entered fully into reproductive
condition. When this animal’s data record is divided
into successive fifths, he will contribute data to blocks
representing the mating period for most snakes. This
situation clearly poses an interpretive problem for
Vincentized data because we can infer that this animal’s
scores will probably underestimate movement being
energized by sexual motivation. The few subjects still
“participating” late in the season may possess special
characteristics that are responsible for their continued
availability, and these characteristics likely will affect
the data being recorded from them, perhaps leading
to exaggerated scores. No solution to this problem is
perfect, and Vincentized data ought to be regarded as
useful with the proviso herein expressed.
Following Vincentizing data, between subjects
ANOVA was conducted to examine the mean number
of meters traveled between successive relocations during
late spring and summer, presented in five successive
Vincentized blocks. The rate of movement between
Vincentized blocks was also assessed by applied a 3 x
5 mixed-effects linear model, treating the three groups
of rattlesnakes as a between-subjects factor and the
five Vincentized blocks as a repeated-measures factor.
Newman-Kuels post-hoc comparisons were conducted
following all ANOVAs with alpha set at 0.05 unless
otherwise indicated.
Straightness-of-path during and after vernal migration in Crotalus viridis
Results
In order to compare straightness-of-path during and
after the vernal migration for each snake we used the
same number of post-migration days as were involved
in the snake’s migration. Mean duration was similar
across the groups (No. Days: F2,16 = 0.05; P > 0.05; No.
Locations: F2,16 = 0.11; P > 0.05; Table 1). Although
males traveled numerically farther than females during
the migration and in the immediate post-migration
period, neither of these differences were significant
(Migration: F2,16 = 1.71; P > 0.05; Post-migration: F2,16
= 0.98; P > 0.05). For each group of snakes, however,
the distance traveled during the migration was greater
than the distance during an identical period afterwards
(Table 1). In fact, of the 19 snakes, only two males
failed to travel farther during the vernal migration than
during the immediately following period of the same
duration (χ2 = 11.84; df = 1; P < 0.01). Accordingly, we
are reasonably confident that the criterion used to define
the end of the migration has ecological validity.
All groups exhibited a reduction in r-values following
the migration. Paired t-tests were applied to these data
(Males: t9 = 3.44; P < 0.01; NG females: t3 = 2.37; P
> 0.05; G females: t4 = 2.10; P < 0.20; All females:
429
t8= 3.14; P < 0.05; Table 2). Hence, males provided
unambiguous evidence for reduction in straightness-of-
path after migration. The reductions in both groups of
females were not significant likely because of the small
sample sizes. In fact, all individuals except for one G
female had a smaller r-value after than during the vernal
migration (χ2 = 5.42; df = 1; P < 0.05). After the two
groups of females were pooled, the t-test was significant
(see above); accordingly we conclude that straightness-
of-path declined in all categories of C. viridis upon
reaching their initial hunting sites. Closely paralleling
these findings was the fact that daily distance traveled
declined in all groups of rattlesnakes between the vernal
migration and the days thereafter (Paired t-test, Males: t9
= 2.31; P < 0.05; Paired t-test, NG females: t3= 2.50; P <
0.10; Paired t-test, G females: t4= 3.49; P < 0.05; Paired
t-test, All females: t8= 4.47; P < 0.01). Accordingly,
distance and straightness-of-path exhibited correlated
change. Mean r-values did not differ significantly among
the three categories of C. viridis during the migration
(F2, 16 = 0.12; P > 0.05), nor did they differ significantly
afterwards (F2, 16 = 0.63; P > 0.05). Therefore, although
straightness-of-path declined when the snakes reached
their hunting grounds, the r-values did not drop to the
level of a “random walk” (Table 2). Mean daily distance

Table 1. Mean duration (days) of the vernal migration, mean numbers of times animals were located during

the migration, mean distance (m) covered during that period, and mean distance covered during the

Table 1. Mean duration (days)

immediate of the vernal
post-migration migration,
period. Numbersmean numbers
in parentheses areof times errors
standard animals were
of the located
respective during the migration, mean
means
distance (m) covered during that period, and mean distance covered during the immediate post-migration period. Numbers in
(SEM). Mean duration and mean number of locations during the post-migration period were the same as
parentheses are standard errors (SE). Mean duration and mean number of locations during the post-migration period were the
same as those of thethose
vernal migration.
of the vernal migration.

________________________________________________________________________
Distance (m)
________________________________________________________________________
Mean Mean no. Migration Post- ta tb
duration locations migration
(days)
_______________________________________________________________________
Males 11.5 (3.6) 4.5 (0.8) 680.3 (114.2) 362.6 (145.2) 2.29* 2.31*
(n = 10)

Gravid 9.6 (4.1) 4.2 (1.5) 463.9 (140.3) 137.6 (105.9) 2.79* 3.49*
Females
(n = 5)

Non-gravid 11.3 (6.6) 4.8 (1.8) 580.2 (402.9) 164.5 (118.5) 1.45 2.50
females
(n = 4)

All females 10.3 (6.2) 4.4 (0.8) 515.6 (181.5) 149.5 (53.4) 2.70* 4.43**
(n = 9)
________________________________________________________________________
ta compares total distance traveled during migration with that traveled during a like period afterwards.
tb compares distance traveled per day during migration with the ratio seen during a like period afterwards.
* p < 0.05; ** p < 0.01; All t-tests reported in this paper are two-tailed tests.
430 David Chiszar et al.

movement (D) did not differ significantly among the
three groups of snakes during the migration (F2, 16 = 1.71;
P > 0.05), nor did they differ in the days immediately
after the migration (F2, 16 = 0.98; P > 0.05). When all
females were combined, the corresponding migration
and post-migration D scores were significantly greater
than zero.
During the remainder of the active season all
snakes were located multiple times, but the number
of locations varied (Table 3), indicating that males
were tracked for fewer days and less frequently than
females.The numerical differences between the three
groups, however, were not statistically significant (No.
Locations: F2, 16 = 0.32; P > 0.05; No. Days: F2, 16 = 1.49;
P > 0.05). Nevertheless, the standard errors (Table 3)
indicate that individual snakes were tracked for quite
different periods of time, the ranges being 13-140 days
(males), 46-146 days (G females), and 47-109 days (NG
females). This creates a statistical problem for measures
of behavior such as straightness-of-path when we plot
means over the late spring and summer. As time passes,
predation, wandering out of range of the receiver, and
battery failure contribute to fewer animals remaining
under observation, and so fewer scores contribute to the
means. For example, by the end of the third month, less
than half of the initial snakes remained detectable (Table
4). Note that more males than females were lost by day
30 (early summer). Chi squares were calculated for the
number of C. viridis remaining under observation for
each 30-day period shown in Table 4, however only the
first 30 day interval showed a signifcance (χ2 = 4.56, df
= 1; P < 0.05). Although losses did not occur during the
vernal migration, they clearly occurred subsequently,
and during the first two months (post- migration) males
were lost more often than females.
Neither distance traveled nor r-value dropped to zero
during the active season (Tables 5 and 6). For most
tests for the movement data, 16 of 20 means were
significantly higher than zero (Table 7). Following
Bonferroni adjustment to compensate for multiple tests
and shifting α to 0.01, nine means remained significantly
higher than zero. For the r-values in Table 6, all 20
means were significantly higher than zero with α =
0.05, and 15 remained significant with α = 0.01. Hence,
the rattlesnakes continued moving through the active
season, sometimes moving only short distances within
an ambushing station (repositioning) and sometimes
shifting from one ambushing station to another.

Table 2. Upper panel: Mean straightness-of-path (r) values and daily distances in meters (D) traveled by
Table 2. Upper panel: Mean straightness-of-path (r) values and daily distances in meters (D) traveled by each group of C. viridis
each group of C. viridis during the vernal migration and during a comparable period of time thereafter.
during the vernal migration and during a comparable period of time thereafter. Lower panel: single-sample t-tests comparing
tabled r-values and Lower
D to zero.
panel: single-sample t-tests comparing tabled r-values and D to zero.

________________________________________________________________________
Migration Post-migration Migration Post-migration
R SEM R SEM D SEM D SEM
________________________________________________________________________
Males .6140 .0901 .3054 .0309 114.4 37.0 32.1 10.3
(n = 10)

Gravid .5221 .1655 .3557 .1535 90.6 38.3 16.7 7.2

Females
(n = 5)

Non-gravid .5361 .1440 .2075 .0621 45.1 13.4 8.1 3.4

Females
(n = 4)

All females .5283 .1052 .2899 .0887 70.4 22.4 12.9 4.3
(n = 9)
________________________________________________________________________
6.81** 9.88** 3.09* 3.11*
3.15* 2.31 2.36 2.38
3.72* 3.34* 3.36* 2.37
5.02* 3.26** 3.14* 3.00*
________________________________________________________________________
* p < 0.05; ** p < 0.01
Straightness-of-path during and after vernal migration in Crotalus viridis
Table 3. Mean number of times snakes were located during late spring and summer, and mean number of
431
Table 3. Mean numberdays C. viridis were tracked during this period (both values exclusive of the figures for locations and days
of times snakes were located during late spring and summer, and mean number of days snakes were
tracked during this shown
periodin(both
Table values
1). exclusive of the figures for locations and days shown in Table 1).

________________________________________________________________________
Mean no. locations SEM Mean no. days SEM
________________________________________________________________________
Males (n = 10) 23.7 4.8 60.0 14.0

Gravid females 29.0 6.0 103.4 20.4

(n = 5)

Non-gravid females 27.2 3.7 79.7 12.9

(n = 4)

All females (n = 9) 28.2 3.5 92.8 12.7

________________________________________________________________________

Gravid females traveled less than either the males or
the NG females (Table 5). Summing over Vincentized
blocks, males traveled a total of 568.2 m, G females
traveled a total of 181.2 m, and NG females traveled
a total of 465.5 m during the late spring and summer
(F2, 16 = 6.98; P < 0.01). The mean for G females was
significantly lower than NG females and males (P = 0.05
and 0.009, respectively) yet these two groups did not
differ significantly from each other (P > 0.05). For both
males and NG females, the rate of movement increased
over Vincentized blocks (Table 5) and the 3 x 5 mixed-
effects linear model revealed a significant effect of
Vincentized blocks (F4, 64 = 31.43; P < 0.01). The mean
for males and NG females on the first Vincentized block
was significantly smaller than the means of all remaining
blocks (all P’s < 0.05). For males, the means on blocks 2-
5 did not differ significantly (P > 0.05); for NG females
the mean on block 4, though significantly higher than
that on block 1, was also significantly lower than the
means on blocks 2, 3, and 5 (P < 0.05). By contrast,
the five means for G females did not differ significantly
except that the mean on block 3 was significantly lower
than that on block 2.
Presented r-values did not differ significantly among
the three categories of snakes (F2, 16 = 0.73; P > 0.05),
nor was there a significant change in these values over
Vincentized blocks (F4, 64 = 1.17; P > 0.05). Hence, the
best estimate of r during the late spring and summer
was the grand mean of 0.37 +0.02, indicating that the
typical movement never represented random direction

Table 4. Number of C. viridis (percent) remaining under observation after various numbers of days in late
Table 4. Percentage
spring C. viridis
of† and summer.under observation after various days in late spring and summer.
†

________________________________________________________________________
30 60 90 120 150
________________________________________________________________________
Males 6 (60%) 5 (50%) 2 (20%) 2 (20%) 0 (0%)
(n = 10)

Gravid 5 (100%) 5 (100%) 3 (60%) 3 (60%) 0 (0%)

Females
(n = 5)

Non-gravid 4 (100%) 3 (75%) 1 (25%) 0 (0%) 0 (0%)

Females
(n = 4)

Sum (19; 100%) 15 (79%) 13 (68%) 6 (32%) 5 (26%) 0 (0%)

________________________________________________________________________
†
Late spring was operationally defined as beginning for each snake at 2days following initial post-surgical
release. In general, this worked out to be the latter half of May.
432 David Chiszar et al.
Table 5. Mean number of meters traveled between successive relocations during late spring and summer,

Table 5. Mean number

presented in five successive Vincentized blocks. Numbers in parentheses are standard errors of the
of meters traveled between successive relocations during late spring and summer, presented in five
successive Vincentized blocks.
respective Numbers
means (SEM). in parentheses are standard errors (SE).

________________________________________________________________________
Vincentized Fifths
________________________________________________________________________
1 2 3 4 5
Males (n = 10) 69.9 (24.9) 133.4 (39.2) 117.7 (26.9) 110.2 (22.1) 136.9 (25.9)

Gravid females 36.2 (9.5) 60.3 (29.0) 15.3 (2.1) 43.0 (24.7) 26.2 (9.7)
(n = 5)

Non-gravid females 18.8 (6.5) 109.6 (26.5) 122.7 (29.9) 68.5 (13.7) 145.8 (27.0)
(n = 4)

All females 28.5 (6.4) 82.2 (20.6) 63.0 (22.4) 54.3 (14.9) 79.3 (24.2)
(n = 9)
________________________________________________________________________

taking. Although 0.37 is certainly not a straight line,
it is far from a random walk. Of the 95 Vincentized
paths (Table 6) (19 snakes x 5 Vincentized blocks =
95), only eight had r-values less than 0.10, far fewer
than we would expect if snakes were taking random
directions at the start of each episode of movement (χ2 =
625.26; df = 1; P < 0.01). Also, 25 of the 95 paths had
r-values in excess of 0.50, and we would expect to see
none of these if movement directions were selected at
random. This last analysis turned up another interesting
point, namely, that 76% of the r’s > 0.50 were seen in
males and 24% were seen in females (χ2 = 4.64; df = 1;
P < 0.05), probably reflecting the fact that males hunt
for prey as well as search for mates, whereas females
only hunt for prey. Mate finding movements likely are
relatively longer and straighter than most prey-related
movements.
In late September and early October, as snakes headed
back to their hibernacula, the paths became straighter
than they were during the late spring and summer.
Although only five snakes were followed during the
autumnal migration (2 males, 3 formerly G females), all
showed larger r-values during this migration than they
did during all or most of the preceeding Vincentized
blocks (Table 8). A repeated-measures ANOVA was
applied to the data of Table 8, revealing a significant

Table 6. Mean r-values

Tableduring
6. Meanlate spring
r-values andlate
during summer, presented
spring and summer, in five successive
presented Vincentized
in five successive blocks.
Vincentized Numbers in parentheses
blocks.
are standard errors Numbers
(SE). in parentheses are standard errors of the respective means (SEM).

________________________________________________________________________
Vincentized Fifths
________________________________________________________________________
1 2 3 4 5
Males .3272 (.06) .4865 (.10) .4795 (.08) .3763 (.05) .3680 (.06)
(n = 10)

Gravid .2527 (.04) .2849 (.06) .3300 (.07) .3692 (.08) .4317 (.07)
Females
(n = 5)

Non-gravid .2634 (.06) .2034 (.05) .2527 (.07) .3882 (.09) .5131 (.16)
females
(n = 4)

All females .2575 (.03) .2487 (.04) .2956 (.05) .3777 (.06) .4676 (.07)
(n = 9)
________________________________________________________________________
Straightness-of-path during and after vernal migration in Crotalus viridis 433
Table 7. Single-sample t-tests comparing means in tables 5 and 6.
Table 7. Single-sample t-tests comparing means in tables 5 and 6 to zero.
________________________________________________________________________
Distance R
Vincentized Fifths Vincentized Fifths
________________________________________________________________________
1 2 3 4 5 1 2 3 4 5

Males 2.80* 3.40** 4.37** 4.98** 5.28** 5.45** 4.86** 5.99** 7.52** 6.13**
(n = 10)

Gravid 3.81* 2.07 7.28** 1.74 2.70 6.31** 4.74** 4.71** 4.61** 6.16**
Females
(n = 5)

Non- 2.89 4.13* 4.10* 5.00* 5.40** 4.39* 4.06* 3.61* 4.31* 3.20*
Gravid
Females
(n = 4)

All
Females 4.45* 3.99** 2.81* 3.64** 3.27** 8.58** 6.21** 5.91** 6.29** 6.68**
(n = 9)
________________________________________________________________________
*p < 0.05 **p < 0.01

effect of Vincentized r-values (F5, 20 = 4.54; P < 0.01).
The mean r-value during the autumnal migration was
significantly higher than the means on the first four of
the five Vincentized blocks. Of course, it must be kept
in mind that this test was conducted a small subset of
our snakes. Nevertheless, we tentatively conclude that
the autumnal migration was characterized by a greater
straightness-of-path than was seen, on average, during
the feeding and mating season. Also, comparing the
right-most column of Table 8 with the migratory r-
values of Table 2, it can be seen that r-values were
slightly higher during the vernal migration than during
the autumnal migration. Each of these snakes eventually
returned to the same C. ludovicianus colony from which
it emerged in the spring (Shipley et al., 2013).

Table 8. r-values on the five Vincentized blocks and during the autumnal migration for 5 C. viridis.
Table 8. r-values on the five Vincentized blocks and during the autumnal migration for 5 C. viridis.
________________________________________________________________________
Vincentized Blocks
________________________________________________________________________
Snake ID 1 2 3 4 5 Autumnal
Migration
________________________________________________________________________
Male 93 .0575 .1966 .0626 .1086 .4308 .5612

Male 517 .2116 .1088 .3408 .1438 .1301 .4891

Female 133 .1280 .1461 .4214 .3020 .3103 .4235

Female 635 .1853 .4878 .3973 .5125 .7001 .6213

Female 893 .3311 .1703 .3257 .0641 .3299 .3766

a a a a a, b
Means .1827 .2219 .3095 .2262 .3802 .4943b

Means with common superscripts do not differ significantly, p > 0.05.

434
Discussion
The main findings of this study are as follows: (1) r-
values during the vernal migration were higher than those
seen at any time later in the active season; (2) although
average r-values and distance moved diminished after
the vernal migration, neither of these dropped to zero; (3)
after a post-migration pause, distance moved increased
for males and NG females, though not to the levels seen
during the vernal migration; and (4) for the five animals
that were tracked during the autumnal migration, r-
values increased during this period over levels during
late spring and summer.
Several points must be kept in view as one interprets
data derived from radio telemetry. Distance traveled
and straightness-of-path are usually underestimated
unless animals are continuously monitored, an almost
impossible requirement for snakes. Thus, our maps will
probably generate lower r-values than a continuous
record would produce. Continuous monitoring over long
periods will eliminate all of these problems, so location
data taken at multi-day intervals must be interpreted
cautiously.
Straightness-of-path during migration can have
several adaptive advantages, such as economy of
effort in getting from the hibernaculum to the hunting
grounds, minimization of exposure to predators during
the trip, and enhanced probability of encountering prey.
It is also possible that straightness-of-path of the vernal
migration can facilitate the return during the autumnal
migration. Since Duvall, King and Gutzwiller, (1985)
saw high r-values in Wyoming samples of C. viridis,
we were not surprised to see relatively high scores in
Colorado. It is interesting, however, that the Wyoming
snakes appear to have much straighter vernal migratory
paths (average r-value = 0.98) than the Colorado
snakes (average r-value = 0.57), and we attribute
this difference to shorter migrations and greater prey
availability at our Colorado site (Shipley et al., 2013).
Pocket gophers (Geomys bursarius Shaw, 1800) were
especially common at PCC wherever soil was disturbed
by C. ludovicianus or by human activities. Furthermore,
long, straight movements have also been documented in
the Desert massasauga (Sistrurus catenatus edwardsii)
where prey availability is at a much lower abundance in
habitats near winter hibernacula (Wastell and Mackessy,
2011). Further, since r-values stayed significantly above
zero, this may be suggestive of most movements being
directed either by external cues (Clark, 2004, 2007) or
by interoceptive ones (e.g., memory). Of course, some
of the longer movements of males may have been
David Chiszar et al.
associated with mate finding, and the relatively high
r-values imply either that males remembered where
females might be found or that the females were detected
from considerable distance such that males could travel
more-or-less straight to them (Ashton, 2003). Also,
males traveling in relatively straight paths may have a
higher probability of intercepting trails made by females
than would be the case for males taking a meandering
course (Duvall et al., 1997; Duvall and Schuett, 1997).
In short, neither males nor females were simply moving
in a random manner at any time during the active
season.
Perhaps the most important aspect of our results was
the fact that our snakes never reduced their paths to
random turnings and meanderings. They moved around
within hunting areas, sometimes in apparently random
ways while taking up a variety of ambushing stations.
Yet, when they traveled between hunting areas, the paths
were again fairly straight, causing their overall r-values
to exceed zero. This same inference is not necessarily
appropriate for the vernal migration, especially in the
Haystack Mountain area, because straight paths are
known to be most efficient for finding dispersed prey,
regardless of the particular direction taken by the
snake (Duvall, King and Gutzwiller, 1985; Duvall et
al., 1997). Hence, natural selection could have favored
individuals who migrate along straight paths without
having a specific direction. During the feeding season
in both the Haystack Mountains and PCC study sites,
however, movements are shorter than those seen during
the vernal migration and they appear to have geographic
goals (e.g., specific hunting areas that these snakes or
other conspecifics have visited before). One reason
this view seems sound is that individual snakes in our
study were seen to move back and forth between several
hunting areas, along the same straight paths. Other
findings provide further support for the general points
that snakes become familiar with their home ranges
and move in a more-or-less directed manner therein:
(1) snakes are reliably found at particular ambush and
refuge locations (Gregory, Macartney and Larsen, 1987;
Reed and Douglas, 2002); (2) often the same paths
are used in moving between these locations (Klauber,
1956; Gannon and Secoy, 1985); (3) sometimes the
same shedding sites are used in successive years
(Parker and Anderson, 2007); and (4) an escape route
may not only be used repeatedly but sometimes the
route is traveled backwards so that the snake’s head
remains focused on the danger, implying that the snake
was probably not attending to landmarks or chemical
cues associated with the escape route (Klauber, 1956).
Straightness-of-path during and after vernal migration in Crotalus viridis
Relatively high r-values during the feeding season
generally imply directed movements and familiarity
with the terrain. However, Mosauer (1933) asserted that
rattlesnakes moved at random during nighttime feeding
forays. Although active movements during nocturnal
hours would not compromise the snakes crypsis, our
results, together with those of the authors cited above,
demonstrate that this assertion seems incorrect (see also
Mosauer, 1935). Indeed, videography of both the red
diamondback rattlesnake (Crotalus ruber) and northern
Pacific rattlesnake (C. oreganus oreganus) in ambush
positions indicated chemosensory probing and mouth
gaping as the two most common behaviors observed
during nocturnal hours (Barbour and Clark, 2012).
Perhaps we ought to be phrasing this speculation in
terms of cognitive maps or similar spatial representational
schemata within the snakes’ nervous systems.
Certainly the r-values point toward the feasibility of
such speculation, yet until proper experimentation
can be conducted, behavior in regards to cognitive
maps should presently be considered as incomplete
(Pearce, 1987). Perhaps a first step in the development
of theory for the causal regulation of r-values in free
ranging snakes is suggested by the idea that PCC snakes
were in better condition than the Wyoming specimens
(Duvall, King and Gutzwiller, 1985) and that PCC
habitat was potentially richer in prey than the Haystack
Mountain habitat (see Shipley et al., 2013, for data).
Experimental translocation of young snakes between
these habitats might reveal whether r-values exhibit
environmentally-induced plasticity. This test would
suggest that upon translocation, Wyoming snakes
would behave like Colorado snakes and vice versa.
Translocation studies sometimes have encountered
difficulties arising from snake disorientation and
associated problems, perhaps because the animals were
moved from familiar to unfamiliar areas. To avoid these
problems, in situ experiments could manipulate feeding
planes and, hence, snake condition, to determine if this
interoceptive factor influences subsequent migratory
behavior. Related experiments could alter density of
prey or density of prey-related cues to assess the extent
to which these exteroceptive factors affect migratory
behavior. At present, no assertion can be made regarding
results of any of these proposed studies, except to say
that such data are needed in order to develop insight into
causal mechanisms underlying the topography of snake
movement. Furthermore, such studies will reveal the
extent to which survival of captive-raised, head-started,
or otherwise provisioned individuals might depend
upon the properties of the release sites. For example,
435
it is entirely possible that snakes in excellent condition
might fail to thrive in marginal habitats with which
locally adapted conspecifics cope effectively.
Acknowledgements. The authors thank the staff of the Plains
Conservation Center for permission to work on the grounds and
for various forms of assistance during the project. We also thank
Alameda East Veterinary Hospital for the use of surgical facilities
and for staff assistance during surgeries. Guidelines for all
necessary animal care were established by a qualified veterinarian
and followed ASIH “Guidelines for Use of Live Amphibians and
Reptiles in Field and Laboratory Research” (http://www.asih.
org/files/hacc-final.pdf). Snake captures were permitted under
Colorado Division of Wildlife Scientific Collection License No.
HP894.
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