CYCHRINE GROUND BEETLE (COLEOPTERA: CARABIDAE: CARABINAE:

CYCHRINI) SURFACE ACTIVITY IN THE NORTHERN COAST RANGE OF

OREGON

By James J. Kirk
150 Jenkins Creek Road
Goldendale, Washington 98620

10 January 2008

CONTENTS

Introduction
Conclusions
Methods
Trapping in 2003
Trapping in 2004 and 2005
Results
Discussion
Occurrence
Onset of Surface Activity
Differences in Mean Trap Dates
Mean Trap Dates
Intraspecific Differences in Mean Trap Dates at Different Sites
Sex-Related Differences
Surface Activity and Precipitation
Surface Activity and Temperature
Soil Temperature
Air Temperature
Acknowledgments
Literature Cited

INTRODUCTION

The purpose of this study was to learn when cychrine beetles (Coleoptera: Carabidae:
Carabinae: Cychrini) are active at the surface in the northern Coast Range of Oregon.

Johnson et al. (1966) pitfall-trapped ground beetles for one year in three habitats in
southwestern Washington. They reported weekly trap results for four species of ground
beetle, including Scaphinotus angusticollis Mannerheim. Pearsall (2003) pitfall-trapped
ground beetles for four or five months in various forest habitats at two sites in British
Columbia, and reported monthly trap results for a variety of ground beetles, including
Scaphinotus angusticollis, S. marginatus (Fisher) and Cychrus tuberculatus Harris (S.
angulatus Harris also was trapped, but too infrequently for quantitative analysis). I am
aware of no reports of multi-year studies focused on northwestern cychrine beetle surface
activity.

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LaBonte (1998) summarized the natural history of Scaphinotus angusticollis and S.
marginatus. He also discussed the pitfall susceptibility of Scaphinotus angusticollis, and
the bias that may arise in comparing pitfall trap results for different species of
Scaphinotus.

CONCLUSIONS

Six cychrine species were trapped: Cycrhus tuberculatus Harris, Scaphinotus angulatus
Harris, S. angusticollis Mannerheim, S. marginatus (Fischer), S. r. rugiceps (Horn), and
S. velutinus Ménétriés.

Scaphinotus angusticollis and S. velutinus were more common near the coast.
Scaphinotus angulatus and S. rugiceps were more common inland; S. rugiceps occurred
only at the most-inland site just east of the Coast Range crest. Scaphinotus marginatus
and Cychrus tuberculatus showed no obvious differences between coastal and inland
sites.

The onset of surface activity in Scaphinotus marginatus was significantly earlier than in
S. angusticollis or Cychrus tuberculatus. With one exception out of 20 comparisons, the
onset of surface activity of Scaphinotus marginatus was consistently earlier than in all
other species. The onset of surface activity in Cychrus tuberculatus was significantly
earlier than in Scaphinotus angusticollis. Other interspecific differences in onset of
surface activity were not statistically significant (due mainly to large variances).

The mean trap dates for Scaphinotus angusticollis were significantly later than the mean
trap dates of all other species except S. rugiceps (which did not occur with S.
angusticollis). Scaphinotus angusticollis showed very limited surface activity in the
spring, with a sudden, huge surge in surface activity in the fall. No other species showed
such a sudden and profound surge in surface activity. A slight majority of all cychrines
collected were Scaphinotus angusticollis. Most of the Scaphinotus angusticollis were
from the two sites closest to the coast, where it far outnumbered all other cychrines. The
differences in mean trap dates suggest the possibility of character displacement: the
surface activity peaks of other cychrines may have shifted in response to competition
with Scaphinotus angusticollis during its huge fall peak.

The significant differences in mean trap dates of Scaphinotus angusticollis and S.

velutinus tends to confirm that they are distinct species, as has been generally accepted
since Gidaspow’s (1973) proposed split based on external morphology.

Mean trap dates for Scaphinotus marginatus were significantly earlier than the mean trap
dates of any other species. This is not particularly surprising, given the much more
northerly distribution of Scaphinotus marginatus.

Intraspecific variation in mean trap dates between sites generally appears not to be
significant. The conservative Tukey test showed significant intraspecific differences only

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for Scaphinotus marginatus and only between two pairs of sites. Because the variances
were not homogeneous, even those results must be viewed with caution. However, there
was a significant correlation between the onset of surface activity and elevation in
Cychrus tuberculatus: the higher the elevation, the later the onset of surface activity.
The best data were available for Cychrus tuberculatus, and correlations between onset of
surface activity and elevation were not significant for Scaphinotus marginatus, S.
angusticollis, and S. velutinus (data for S. angulatus and S. rugiceps were insufficient for
analysis).

Females tend to become active on the surface earlier than males. In 13 of 16

comparisons, the mean trap dates of females were earlier than those of males.

There was no significant correlation between trap success and precipitation. Inspection
suggests that the onset of fall rains and the onset of surface activity in Scaphinotus
angustiollic may coincide, and that peaks in precipitation may coincide with peaks in
surface activity of S. marginatus.

Soil temperatures at the onset of surface activity in Scaphinotus marginatus and S.

angulatus were significantly lower than those in S. angusticollis and S. velutinus. Soil
temperatures at the onset of surface activity in Cychrus tuberculatus were significantly
lower than those in Scaphinotus angusticollis. In all of these cases the lower soil
temperature appears to be associated with the earlier onset of surface activity.

Air temperature at the onset of surface activity in Scaphinotus marginatus differed

significantly from air temperature at the onset of surface activity of all other species,
which would be expected given the earlier onset of surface activity in S. marginatus.
Differences among other species in air temperature at onset of surface activity were not
significant.

METHODS

I carried out preliminary pitfall trapping during 2003 at the Hamlet site (see Table 1).
This site was selected because it was on property my wife and I own, and because I had
seen large numbers of Scaphinotus angusticollis active during the day in years past.

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 Table 1: Trap sites.

SITE COUNTY LATITUDE LONGITUDE ELEV DESCRIPTION

(FEET)
East-facing canyon side above the Little
North Fork of the Nehalem River, with
Hamlet Clatsop 45º 51.3’ N 123º 41.8’ W 900 mature alder dominant and some cedar,
spruce, hemlock, and Douglas fir present.
The site has been disturbed by past
logging (ring count of a large spruce cut
in 2004 suggests the last logging may
have been about 80 years ago), and
current use associated with a vacation
home. The area reportedly was used for
dairy farming in the first half of the 19th
century.
Northeast of turnout on north side of Cape
Lookout Road, 1.35 miles via Cape
Cape Tillamook 45º 20.00’N 123º 57.12’W 430 Lookout Road southeasterly from the
Lookout turnoff for the Cape Lookout trailhead,
2.00 miles via Cape Lookout Road
northwesterly from the junction of Cape
Lookout road and Sandlake road. South-
facing hillside with dense alder and
hemlock forest. Very little underbrush.
1.1 miles up gravel road (USFS 1106)
from intersection with Sand Lake Road at
Sand Lake Tillamook 45º 18.43’N 123º 53.22’W 610 a point 1.7 mi east of the junction of Cape
Road Lookout Road and Sand Lake Road.
Gently sloping ridge. Dense, older second
growth hemlock with very little
underbrush.
East of Niagara Road (8533),
approximately 2.45 miles via Niagara
Niagara Tillamook 45º 13.80’N 123º 37.95’W 1,450 Road southerly from the junction with the
Road Nestucca River Road, Tillamook County,
Oregon; 45º 13.80’N, 123º 37.95’W;
elevation 1450 feet. Gently sloping ridge.
Dense alder, with some hemlock and
Douglas fir, vine maple, and sword fern.
Tillamook 45º 16.96’N 123º 31.76’W 1,000 Just north of the Nestucca River Road,
just west of BLM Road 3-7-32, east of an
Nestucca unnamed tributary to the Nestucca River.
Road Moderately sloping area adjacent to a
tributary to the Nestucca River. Mixed

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 alder and Douglas fir, with vine maple
and salmonberry undergrowth..
Yamhill 45º 18.14’N 123º 22.71’W 1,690 South of Fall Creek Road (BLM Road 3-
6-24), just west of its intersection with the
Fall Creek Nestucca Access Road, about 1.75 miles
Road southeasterly via the Nestucca Access
Road from the dam at McGuire Reservoir.
Steep, south-facing hillside, with dense
second-growth conifer forest, with
scattered stands of Oregon grape and
salal, and with scattered sword ferns and
very little underbrush.

Trapping in 2003

I made the pitfalls from plastic deli tubs (Anchor Packaging, D16WX). Each tub was in
the shape of an inverted, truncated cone. The larger plane of the truncated cone (i.e.,
mouth of the deli tub) was open, with an inside diameter of 11 cm. The smaller plane of
the truncated cone (i.e., bottom of the deli tub) was closed, 9 cm in diameter, and 8 cm
along the axis of the cone below the larger plane. The sides of the cone were about seven
degrees from parallel with the axis of the cone, or flared out about seven degrees from
vertical.

I positioned the deli tubs in the soil with their lips at or below the normal soil level, and
set them in an irregular line at each site between 5 and 80 meters apart. I placed a 30-cm
square piece of 30-pound roll roofing over each trap, and supported the roofing about 1
cm off the ground on three or four sticks or twigs pushed vertically into the ground
between the trap and where the edge of the roofing would fall. Undiluted Prestone
LowTox (automotive) Antifreeze/Coolant was placed in each tub to a depth of about 3 to
4 cm.

During 2003 at the Hamlet site, I left the traps open for approximately two weeks at a
time, and closed them with deli tub lids for approximately two weeks between each
trapping period. I replenished the antifreeze as necessary. I collected trap contents at the
end of each trapping period by dumping each trap into a second deli cup. I stored the trap
contents in the antifreeze from the trap, and sometimes added isopropanol.

Trapping in 2004 and 2005

My experience during 2003 led to several modifications used throughout the remainder of
the study. I abandoned roll roofing for trap shelters because it was prone to being
displaced by wind or animals, and tended to droop when heated by the sun. Instead I
used inexpensive, 29-cm square, Saltillo handmade tiles that were approximately 1.5 cm
thick. The vertical, supporting sticks had to be stouter because of the greater weight of a

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tile. I sometimes substituted a pebble, fir cone, or short length of wood laid on the
ground in place of a stick pushed into the ground to support a tile. I made the traps by
nesting one deli tub inside another, so that the lower or outer tub remained in the ground
as a sleeve to keep the soil in place, and the upper or inner one could be removed to
retrieve trap contents. I collected trap contents by dumping the trap through a culinary
screen-type strainer and into a second deli tub. The contents of the strainer were then
placed in a labeled deli tub in 70- to 80-percent isopropanol or ethanol for storage. The
strained antifreeze was returned to the trap. My experience during 2003 also led me to
conclude that continuous trapping with checks every two weeks was necessary to ensure
all surface activity was detected and to improve resolution of surface activity events. I
replaced antifreeze when it became highly diluted with water, which apparently resulted
from condensation, precipitation when tiles were displaced, and possibly hygroscopy.

I removed all of the traps from the Hamlet site in 2003. I placed the traps at the other
sites (see Table 1) in January and February of 2004, and kept them in use through
December 2005. The 2004-2005 sites were selected in an attempt to sample a roughly
east-west transect from the coast to the crest of the Coast Range. Specific sites were
selected based on my assessment of suitable habitat, convenience of access, and low
probability of human disturbance. I tried to check all traps at approximately two-week
intervals, but, because this was an avocational endeavor, the constraints of job and family
often interfered with the precision of that schedule.

On the rare occasions when traps were disturbed by animals or humans, any data from the
disturbed traps were excluded.

In October of 2004, I installed improvised rain gauges at each site. I made each gauge
from a 31-centimeter (12 inches) length of thick-wall (7 millimeter) black plastic pipe
with an inside diameter of 10 centimeters (4 inches). I cemented a standard plastic cap
on one end. I placed each gauge by burying the capped end in about 5 centimeters of soil
so that the cylinder would remain vertical with the opening at the top. The trap-site rain
gauges were situated under the forest canopy, and received less precipitation than if they
had been situated in the open. I placed one additional rain gauge in the open near the
paved road at the Cape Lookout site. Rainfall was measured in the field using a ruler or
tape measure, and the water was then poured into a jug, capped, and taken to the lab to be
weighed. The gravimetric measurement was used in calculations, except that the field
measurement by ruler or tape measure was used where a gravimetric measurement was
not available (e.g., on the rare occasions when the water was frozen or the water was
spilled). Except when the water was frozen, the gauge was emptied after each
measurement. On a few occasions a gauge was tipped by wind or animals – in those
cases a measurement was estimated by interpolating based on nearby gauge
measurements.

In October of 2004, I also started measuring the temperature of the top centimeter or two
of soil at each trap site. I used a calibrated alcohol thermometer and allowed it to
equilibrate while I checked the traps.

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I pinned and labeled all adult cychrines, and identified them using Gidaspow’s (1968,
1973) keys and descriptions. I stored all larval cychrines in capped glass vials in 80
percent ethanol. All other trapped animals were stored in covered deli tubs in 70-percent
isopropanol or 80-percent ethanol. All specimens remain in my collection at the time I
write this.

RESULTS

The total numbers of adults of each species trapped at each site are shown in Table 2.
The overall trap success for each species at each site is summarized in Table 3. Graphic
summaries of surface activity are displayed in Appendix Figures 1 through 19.

TABLE 2: Numbers of Cychrines Trapped at Sites in the Northern Coast Range of

Oregon. Trapping at the Hamlet site was in 2003; trapping at the other sites was in 2004
and 2005.

SITE CAPE SAND FALL

HAMLET LOOKOUT LAKE NIAGARA NESTUCCA CREEK TOTALS
SPECIES
Scaphinotus 51 28 1 60 19 11 170
marginatus
Scaphinotus 1 3 13 17
angulatus
Cychrus 11 21 35 29 24 15 135
tuberculatus
Scaphinotus 37 60 1 2 3 103
velutinus
Scaphinotus 57 300 133 26 8 524
angusticollis
Scaphinotus 57 57
rugiceps
TOTALS 120 386 229 119 53 99 1006

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TABLE 3: Number of Cychrines Pitfall-Trapped Per 100 Trap Nights at Sites in
the Northern Coast Range of Oregon. Trapping at the Hamlet was site was in 2003;
trapping at the other sites was in 2004 and 2005. The “totals” by species in the far right
column were calculated using total numbers of beetles and total numbers of trap-nights;
and do not equal the means of the rows because of site-to-site variation in numbers of
trap-nights (caused by disturbed traps, more traps set at Hamlet, etc.

SITE CAPE SAND FALL

HAMLET LOOKOUT LAKE NIAGARA NESTUCCA CREEK TOTAL
SPECIES
Cychrus 1.15 .61 1.03 .84 .70 .44 .75
tuberculatus
Scaphinotus .10 0 0 .09 0 .38 .09
angulatus
Scaphinotus 5.94 8.77 3.90 .76 .23 0 2.90
angusticollis
Scaphinotus 5.31 .82 0 1.75 .56 .32 .94
marginatus
Scaphinotus 0 1.08 1.76 .03 .06 .09 .57
velutinus
Scaphinotus 0 0 0 0 0 1.67 .32
rugiceps
TOTALS 12.50 11.28 6.69 3.47 1.55 2.90

DISCUSSION

Occurrence

Species-specific differences in pitfall trap susceptibility were not investigated as part of

this study, and apparently have not been reported elsewhere (however, see La Bonte's
(1998) discussion of pitfall trap susceptibility of Scaphinotus angusticollis). Because the
differences in species-specific pitfall trap susceptibility are not known, the interspecific
differences in numbers of beetles trapped and trap success are not reliable indicators of
interspecific differences in population density or magnitude of surface activity. They
probably are reasonably reliable indicators of relative differences in population density
and magnitude of surface activity for the same species at different sites. So, for example,
it would be correct to infer from the tables that the population density, or at least the
magnitude of surface activity, of Scaphinotus angusticollis was greater at the Cape
Lookout site than at the Nestucca site. But it would be incorrect to infer, for example,
that the population density or magnitude of surface activity of Scaphinotus angusticollis
was greater than that of Cychrus tuberculatus at the Cape Lookout site.

The raw numbers of beetles trapped at each site (Table 2) are of interest, but trap success
(Table 3) provides a more accurate indication of relative surface activities because it
takes into account differences in trapping intensity. In a few cases, traps were disturbed,
and the temporary absence of those traps is reflected in trap success. Also, trapping

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intensity was higher at the Hamlet site during 2003 than at the other sites, so comparison
of raw numbers would be misleading.

The frequency data from 2004 and 2005 (Table 2) reveal some patterns. Scaphinotus
angusticollis and S. velutinus were more common near the coast, and less common
inland. Scaphinotus angulatus and S. rugiceps were more common inland, and not found
near the coast. Scaphinotus marginatus and Cychrus tuberculatus show neither pattern,
and seemed to be as common near the coast as inland. The 2003 data from Hamlet were
consistent with those patterns.

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 Onset of Surface Activity

The date a cychrine species first appears in pitfall traps marks the detectable onset of
surface activity for that species. The first dates of appearance are presented in Table 4.

Table 4. Earliest Pitfall Trap Dates of Cychrines in the Northern Coast Range of
Oregon. Gregorian date / Julian date. Asterisks indicate the data are not sufficient to
reliably detect the onset of activity (total samples sizes of one or a few individuals).
Dates are arbitrarily set at the midpoints of pitfall trapping periods. Site-specific
comparisons of earliest dates of appearance are presented in Table 5.

2003
HAMLET CAPE SAND LAKE NIAGARA NESTUCCA FALL CREEK
SPECIES LOOKOUT

S. marginatus Mar 8 / 67 No trapping No trapping No trapping No trapping No trapping

S. angulatus Sep 10 / 253* No trapping No trapping No trapping No trapping No trapping
C. tuberculatus Apr 22 / 112 No trapping No trapping No trapping No trapping No trapping
S. velutinus -- No trapping No trapping No trapping No trapping No trapping
S. angusticollis Apr 22 / 112 No trapping No trapping No trapping No trapping No trapping
S. rugiceps -- No trapping No trapping No trapping No trapping No trapping

2004
HAMLET CAPE SAND LAKE NIAGARA NESTUCCA FALL CREEK
SPECIES LOOKOUT

S. marginatus No trapping Feb 23 / 55 Oct 30 / 304* Feb 23 / 55 Apr 4 / 95 Apr 18 / 109

S. angulatus No trapping -- -- Oct 30 / 303* -- May 3 / 124
C. tuberculatus No trapping Apr 18 / 109 Apr 4 / 95 May 3 / 124 May 18 / 139 Jun 12 / 164
S. velutinus No trapping Apr 18 / 109 May 3 / 124 Oct 2 / 275* Aug 21 / 234* Apr 4 / 95
angusticollis No trapping May 3 / 124 Apr 18 / 109 May 30 / 151 Sep 5 / 249 --
rugiceps No trapping -- -- -- -- May 3 / 124

2005
HAMLET CAPE SAND LAKE NIAGARA NESTUCCA FALL CREEK
SPECIES LOOKOUT

S. marginatus No trapping Jan 13 / 13 -- Mar 27 / 86 Mar 14 / 73 --

S. angulatus No trapping -- -- Apr 23 / 113 -- Mar 27 / 86
C. tuberculatus No trapping Apr 23 / 113 Apr 23 / 113 Apr 23 / 113 Apr 23 / 113 May 21 / 141
S. velutinus No trapping May 21 / 141 May 6 / 126 -- -- --
S. angusticollis No trapping May 21 / 141 May 21 / 141 Apr 23 / 113 -- --
S. rugiceps No trapping -- -- -- -- Apr 23 / 113

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Table 5. Interspecific Differences in Earliest Appearance Dates for Cychrines in the
Northern Coast Range of Oregon. Each number is the result of subtracting the earliest
date for the species at the top of the column in which the number appears from the
earliest date for the species at the left of the row in which the number appears at the site
indicated in the second column. Negative numbers are in parentheses. The sites are
shown at the left end of the rows. The year is shown at the top of each column.

I tested the significance of differences in dates of first appearance of the six species. The
results are presented in Table 6. The lack of significance for some pairs may relate more
to the small sample sizes than to lack of real differences.

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Table 6. Significant Differences in Earliest Dates of Surface Activity of Cychrines in
The Northern Coast Range of Oregon, 2003 To 2005. The “Differences” are the
number of days between the earliest dates of appearance of the two species listed in the
first column at all sites where the two species occur together. Significance was tested
using the t-test. Non-significant differences and comparisons with too-small samples
sizes are not shown.

SPECIES DIFFERENCES (DAYS) n MEAN SIGNIFICANT

AT p = .10?
S. marginatus before 27, 40, 44, 45, 54, 55, 69, 8 54 yes
C. tuberculatus 100
S. marginatus before 27, 45, 69, 96, 128, 154 6 86 yes
S. angusticollis
C. tuberculatus before 0, 0, 14, 15, 27, 28, 28, 110 8 28 yes
S. angusticollis

I tested the correlation between date of first appearance and elevation for C. tuberculatus,
S. marginatus, S. angusticollis, and S. velutinus. Insufficient data were available to
calculate meaningful correlations for the other species. The correlation was significant
(t-test, p = .05) only in C. tuberculatus, which is the species with the best data. See
Figure 1.

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Figure 1. Elevation and Date of First Appearance of Cychrus tuberculatus in the
Northern Oregon Coast Range. R = 0.75, b = 18.2, y = -1178 + 18.2x, t = 3.359

Differences in Mean Trap Dates

I trapped the Hamlet site during 2003 and the other five sites during 2004 and 2005, for a
total of 11 trap-site-years. Two to four species occurred together during each trap-site
year, yielding a total of 36 species data sets containing 8 or more individual trap dates (I
concluded that data sets with fewer than 8 trap dates would not provide reliable estimates
of mean trap dates). I calculated the mean trap dates for each of those 36 data sets. I then
calculated the difference in mean trap dates for each species pair for each trap-site-year. I
aggregated the differences in mean trap dates from all trap-site-years for each of the six
species pairs for which there were two or more sets of means, and performed t-tests with
a null hypothesis that the true difference between the means was zero. The results appear
in Table 7.

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Table 7. Differences in Mean Trap Dates. Differences in mean trap dates for species
pairs from all sites and all years. Abbreviations have the following meanings: angl=
Scaphinotus angulatus; angu = S. angusticollis; marg = S. marginatus; rug = S.
rugiceps; tub = Cychrus tuberculatus; vel = S. velutinus. The two-tailed probabilities of
obtaining the observed result if the null hypothesis is true are from Dr. Robert Knodt’s
MODSTAT.

angl-rug angu-mar angu-tub angu-vel tub-marg vel-tub

0 141 113 41 28 71
70 108 35 83 73 51
150 94 65 56 7
83 77 66 11 73
90 6
116 92
139

35.0 120.5 94.9 63.8 44.3 50.8 means

2450 951 1108 298.2 1215 953.4 variance
1.00 7.82 7.53 7.38 3.10 3.30 t calc
12.71 3.182 2.447 3.182 2.571 3.182 t-tab .05
0.500 0.0036 0.0006 0.0042 0.0267 0.0441 p 2-tailed

I draw the following conclusions from the analysis of the differences in mean trap dates:
(1) The mean trap dates for S. angusticollis were significantly later than those for
S. marginatus, S. velutinus, and C. tuberculatus;
(2) The mean trap dates for C. tuberculatus were significantly later than those for
S. marginatus;
(3) The mean trap dates for S. velutinus were significantly later than those for C.
tuberculatus; and
(4) The differences between mean trap dates for S. angulatus and S. rugiceps
were not significant.

Mean Trap Dates

In addition to analyzing the differences in mean trap dates, I directly compared

trap dates for all species at all sites in all years using analysis of variance (ANOVA). I
used two ANOVA programs: (1) Dr. Robert Knodt’s MODSTAT software; and (2)
online ANOVA available at www.physics.csbsju.edu/stats/anova.html. MODSTAT
warned that variances were not homogenous (Bartlett’s test), and thus that the test may
not be valid. Because the sample sizes varied widely (17 for S. angulatus to 524 for S.

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angusticollis), and the variances were not homogeneous, the probability of a Type I error
was substantially increased. Elimination of S. angulatus from the analysis did not make
the variances homogeneous, so I kept S. angulatus in the analysis and randomly (using a
random number table) drew a sample from the trap dates for each of the other species so
that all sample sizes were equal (sample size for each was 17). MODSTAT then reported
that the variances were homogeneous (Bartlett’s test and Hartley’s test). The ultimate
results of those two MODSTAT tests were the same. I infer that the online ANOVA
discarded outliers until the variances were homogeneous; nevertheless, I conducted the
online ANOVA both with all data and with the randomly selected subsets. The results of
these four ANOVAs are summarized in Table 8.

Table 8. Summary of Four Analyses of Variance Tests Run on Trap Dates.

Dr. Robert Knodt’s MODSTAT ANOVA, all data

Source of Mean Probability
Variation Df Sums of Squares Squares F (Null Hypo)
Between 5 3454279.5308 690855.9062 202.0462 0.0001
Within 1000 3419296.3600 3419.2964 ---
Total 1005 6873575.8908 --- ---

Dr. Robert Knodt’s MODSTAT ANOVA, randomly selected subsets for equal samples
sizes of 17
Source of Mean Probability
Variation Df Sums of Squares Squares F (Null Hypo)
Between 5 256054.0000 51210.8000 12.0352 0.0001
Within 96 408486.8800 4255.0717 ---
Total 101 664540.8800 --- ---

Online ANOVA (www.physics.csbsju.edu/stats/anova.html), all data

Source of Mean Probability
Variation Df Sums of Squares Squares F (Null Hypo)
Between 5 3461400 692270 202.4 0.0001
Within *** 3419700 3420
Total *** 6881100

Online ANOVA (www.physics.csbsju.edu/stats/anova.html), randomly selected subsets

for equal samples sizes of 17
Source of Mean Probability
Variation Df Sum of Squares Squares F (Null Hypo)
Between 5 256560 51313 12.06 0.0001
Within 96 408330 4253
Total 101 664890

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MODSTAT compared means using the t-test, the Duncan Multiple Range Test, and the
Tukey test. The results are displayed in Table 9.

Table 9. Interspecific MODSTAT Comparisons of Trap Dates.

A test result displayed before the slash is for all data (variances not homogeneous); a test
result displayed after the slash is for the randomly selected subsets to give equal sample
sizes of 17 (variances homogeneous). Significant (.05) t values are in boldface.

Comparing With Tukey test

t-test Duncan test Crit val = 57/66

S. angulatus S. angusticollis 5.965 / 3.174 Sig / Sig 71 / 72

S. angulatus C. tuberculatus 3.082 / 0.606 Sig / -- --
S. angulatus S. marginatus 3.649 / 3.208 Sig / Sig 79 / 88
S. angulatus S. rugiceps 1.479 / 1.784 Sig / -- -- / --
S. angulatus S. velutinus 0.259 / 0.692 -- / -- -- / --
S. angusticollis C. tuberculatus 24.188 / 4.967 Sig / Sig 105 / 86
S. angusticollis S. marginatus 28.895 / 7.264 Sig / Sig 150 / 160
S. angusticollis S. rugiceps 6.491 / 1.677 Sig / -- -- / --
S. angusticollis S. velutinus 11.766 / 2.567 Sig / Sig 66 / --
C. tuberculatus S. marginatus 5.718 / 3.294 -- / Sig -- / 74
C. tuberculatus S. rugiceps 8.566 / 3.039 Sig / Sig 61 / --
C. tuberculatus S. velutinus 5.430 / 1.489 Sig / -- -- / --
S. marginatus S. rugiceps 8.625 / 5.674 Sig / Sig 106 / 130
S. marginatus S. velutinus 8.310 / 4.166 -- / Sig 84 / 106
S. rugiceps S. velutinus 1.934 / 1.093 Sig / -- -- / --

The fact that variances are not homogeneous does not necessarily mean ANOVA results
are not trustworthy – the lack of homogeneous variances merely increases the probability
of a Type I error. Since ANOVA results based on heterogeneous variances have some
value, I scored results as follows: (1) a significant t-test, Duncan test, or Tukey test based
on data sets with heterogeneous variances was assigned one point; (2) a significant t-test,
Duncan test, or Tukey test based on data sets with homogeneous variances was assigned
two points; and (3) a significant difference in means analysis from the previous section
was assigned two points. I scored the fifteen species pairs accordingly. The results
appear in Table 10.

Page 16 of 32 -- Cychrine Surface Activity by Kirk

Table 10. Composite Scores Reflecting Significance of Differences in Trap Dates.
See text for explanation.

Comparing With Composite Score

S. angusticollis C. tuberculatus 11
S. angusticollis S. marginatus 11
S. angulatus S. angusticollis 9
S angulatus S. marginatus 9
S. angusticollis S. velutinus 9
C. tuberculatus S. marginatus 9
S. marginatus S. rugiceps 9
C. tuberculatus S. rugiceps 7
S. marginatus S. velutinus 7
C. tuberculatus S. velutinus 4
S. angulatus C. tuberculatus 2
S. angulatus S. rugiceps 2
S. angusticollis S. rugiceps 2
S. rugiceps S. velutinus 2
S. angulatus S. velutinus 0

Perhaps the most fascinating relationship revealed by this analysis is that all of the
species that occur with S. angusticollis have mean trap dates that are significantly
different from that of S. angusticollis. The only species with mean trap dates not
significantly different from S. angusticollis is S. rugiceps, and those two species did not
occur together at any of the study sites. This suggests the possibility of character
displacement due to interspecific competition: species sympatric with S. angusticollis
may be evolving toward surface activity periods that minimize exposure to the huge
irruption of S. angusticollis in late summer and fall.

The significant difference between mean trap dates for S. angusticollis and S. velutinus
also is noteworthy. These two species appear to be closely related, and were considered
conspecific in the past. This difference in phenology supports the taxonomic distinction.

Intraspecific Differences in Mean Trap Dates at Different Sites

Scaphinotus angusticollis

Scaphinotus angusticollis occurred at five of the six sites, but the sample sizes varied
from 8 at the Nestuccca Road Site to 300 at the Cape Lookout site, and variances were
not homogeneous. See Table 11 for the mean trap dates and total numbers of beetles

Page 17 of 32 -- Cychrine Surface Activity by Kirk

trapped. ANOVA results for the 2004 and 2005 trap dates are presented in Tables 12 and
13.

Table 11. Mean Trap Dates and Total Numbers of Beetles Trapped for Scaphinotus
angusticollis at Different Trap Sites. Numbers are Julian dates. Calculations were
done using Dr. Robert Knodt’s MODSTAT software.

SITE MEAN NUMBER 95% CONF RANGE SAMPLE

JULIAN OF STANDARD
DATES BEETLES DEVIATION

Cape Lookout 302 300 297 – 307 133 – 343 30.2

Sand Lake Rd. 279 133 272 – 286 119 – 325 42.6
Niagara Rd. 269 26 253 – 285 119 – 325 40.8
Nestucca Rd. 265 8 237 – 293 256 – 312 17.2
Hamlet 225 57 214 – 236 116 – 299 70.1

Table 12. ANOVA Results – Between-Site (excluding Hamlet) Variation in Mean

Trap Dates of Scaphinotus angusticollis. Bartlett’s Test of Homogeneity indicates a
lack of variance homogeneity (Bartlett value of 25.232 versus Chi-square value for five
groups of 7.815), so these results must be viewed with caution. Based on MODSTAT
results.

Source Df Sum Squares Mean Square F Probability

Between 3 71616 23872 19.669 0.0001
Within 463 561946 1214 ---
Total 466 633561 --- ---

Table 13. Significant Between-Site (excluding Hamlet) Differences in Mean Trap

Dates for Scaphinotus angusticollis. Based on MODSTAT results.

Comparing With Tukey test

t-test Duncan test Crit val =
Cape Lookout Sand Lake 6.379 --- ---
Cape Lookout Niagara 5.154 Significant ---
Cape Lookout Nestucca 3.434 Significant ---
Sand Lake Nestucca 0.918 Significant ---

The trend suggested by the progressive decline in numbers of beetles trapped (see Table
11) moving from the Cape Lookout site, site-by-site, inland to the Nestucca Road Site
(the inland-most site at which S. angusticollis was trapped) is perhaps obvious, but the
correlations of number of beetles trapped with longitude (r =.9110), elevation (r = .8331),

Page 18 of 32 -- Cychrine Surface Activity by Kirk

2005 annual precipitation (r = .6728), and 2005 precipitation during fall surface activity
(r = .5054) are not statistically significant (probably because of the small sample size and
limited numbers of degrees of freedom; with two degrees of freedom, r does not become
significant at the .05 level until it equals or exceeds .95). Nevertheless, based in part on
the ANOVA results, I think the trend probably is real. The reasons for the trend are not
clear, and probably will not become clear without a careful analysis of both biotic and
abiotic factors.

Scaphinotus marginatus

The mean trap dates and total number of beetles trapped at each site are presented in
Table 14. The ANOVA results are presented in Tables 15 and 16. The conservative
Tukey test shows significant differences between two pairs of sites. No trend is apparent
and the reasons for the significant differences are obscure.

Table 14. Mean Trap Dates and Total Numbers of Beetles Trapped for Scaphinotus
marginatus at Different Trap Sites. Numbers are Julian dates. Calculations were done
using Dr. Robert Knodt’s MODSTAT software.

SITE MEAN NUMBER 95% CONF RANGE SAMPLE

JULIAN OF INTERVAL STANDARD
DATE BEETLES DEVIATION

Cape Lookout 146 28 116 - 176 21 – 353 105

Niagara Rd. 125 60 104 - 146 60 – 325 81.3
Nestucca Rd. 144 19 108 - 180 79 – 312 82.7
Fall Cr. Rd. 237 11 190 - 284 116 – 283 58.6
Hamlet 113 51 91 - 135 74 – 299 60.4

Table 15. ANOVA Results – Between-Site Variation in Mean Trap Dates of

Scaphinotus marginatus. Bartlett’s Test of Homogeneity indicates a lack of variance
homogeneity (Bartlett value of 13.997 versus Chi-square value for five groups of 9.488),
so these results must be viewed with caution. Based on MODSTAT results.

Source Df Sum Squares Mean Square F Probability

Between 4 149968 37492 5.9841 0.0003
Within 164 1027502 6265 ---
Total 168 1177470 --- ---

Page 19 of 32 -- Cychrine Surface Activity by Kirk

Table 16. Significant Between-Site Differences in Mean Trap Dates for Scaphinotus
marginatus. Based on MODSTAT results.

Comparing With Tukey test

t-test Duncan test Crit val = 93.55
Cape Lookout Fall Creek 2.700 Significant ---
Fall Creek Hamlet 6.206 Significant 124
Fall Creek Nestucca 3.274 Significant ---
Fall Creek Niagara 4.355 Significant 112

Cychrus tuberculatus

Mean trap dates and number of beetles trapped are presented in Table 17. ANOVA
results are presented in Table 18. The ANOVA results indicate no significant between-
site differences in mean trap dates. The t-test and Duncan test indicate significant
differences between the Niagara and Fall Creek sites (t = 3.554), and between the Sand
Lake and Fall Creek sites (t = 2.279). However, the more conservative Tukey test
indicates no significant differences, as do the Scheffe Procedure and the Newman-Keuls
test.

Table 17. Mean Trap Dates and Total Numbers of Beetles Trapped for Cychrus
tuberculatus at Different Trap Sites. Numbers are Julian dates. Calculations were
done using Dr. Robert Knodt’s MODSTAT software.

SITE MEAN NUMBER 95% CONF RANGE SAMPLE

JULIAN OF INTERVAL STANDARD
DATE BEETLES DEVIATION

Cape Lookout 187 21 172 – 203 116 - 259 39.9

Sand Lake Rd. 172 35 160 – 184 102 - 256 39.3
Niagara Rd. 168 29 154 – 181 119 - 227 26.8
Nestucca Rd. 185 24 170 – 199 119 – 242 29.3
Fall Creek Rd. 197 15 179 – 216 149 - 219 24.5
Hamlet 189 11 168 – 210 116 – 257 52.4

Page 20 of 32 -- Cychrine Surface Activity by Kirk

Table 18. ANOVA Results – Between-Site Variation in Mean Trap Dates of Cychrus
tuberculatus. Bartlett’s Test of Homogeneity indicates a lack of variance homogeneity
(Bartlett value of 23.28 versus Chi-square value for six groups of 11.07), so these results
must be viewed with caution. Based on MODSTAT results.

Source Df Sum Squares Mean Square F Probability

Between 5 13394 2679 2.1591 0.0618
Within 129 160053 1241 ---
Total 134 172448 --- ---

Scaphinotus velutinus

Mean trap dates and number of beetles trapped are presented in Table 19. ANOVA
results are presented in Table 20. ANOVA, the t-test, and the Duncan test suggest the
existence of significant differences between all three sites (Table 21) where S. velutinus
was trapped. However, variances are heterogeneous and the more conservative Tukey
test shows no significant differences.

Table 19. Mean Trap Dates and Total Numbers of Beetles Trapped for Scaphinotus
velutinus at Different Trap Sites. Numbers are Julian dates. Calculations were done
using Dr. Robert Knodt’s MODSTAT software.

SITE MEAN NUMBER 95% CONF RANGE SAMPLE

JULIAN OF INTERVAL STANDARD
DATE BEETLES DEVIATION

Cape Lookout 246 37 223 – 269 116 – 343 66.9

Fall Cr. Rd. 143 3 63 – 223 102 – 171 36.5
Sand Lake 203 60 185 – 221 116 – 325 71.7

Table 20. ANOVA Results – Between-Site Variation in Mean Trap Dates of

Scaphinotus marginatus. Bartlett’s Test of Homogeneity indicates a lack of variance
homogeneity (Bartlett value of -23.455 versus Chi-square value for three groups of
5.991), so these results must be viewed with caution. Based on MODSTAT results.

Source Df Sum Squares Mean Square F Probability

Between 2 59304 29652 6.1577 0.0034
Within 97 467099 4815 ---
Total 99 526403 --- ---

Page 21 of 32 -- Cychrine Surface Activity by Kirk

Table 21. Significant Between-Site Differences in Mean Trap Dates for Scaphinotus
marginatus. Based on MODSTAT results.

Comparing With Tukey test

t-test Duncan test Crit val = 136.2
Cape Lookout Fall Creek 2.614 Significant ---
Cape Lookout Sand Lake 2.942 Significant ---
Fall Creek Sand Lake 1.432 Signficant ---

Scaphinotus angulatus

Scaphinotus angulatus was trapped in sufficient numbers for analysis at only two sites
(Niagara and Fall Creek), and ANOVA shows no significant difference between those
two sites. Mean trap dates and number of beetles trapped are presented in Table 22.
ANOVA results are presented in Table 23

Table 22. Mean Trap Dates and Total Numbers of Beetles Trapped for Scaphinotus
angulatus at Different Trap Sites. Numbers are Julian dates. Calculations were done
using online software at www.physics.csbsju.edu/stats/anova.html.

SITE MEAN NUMBER 95% CONF RANGE SAMPLE

JULIAN OF INTERVAL STANDARD
DATE BEETLES DEVIATION

Fall Creek 202 13 153 – 251 93 – 283 74.8

Niagara. 252 3 150 – 354 119 – 325 115
Hamlet 257 1 --- --- ---

Table 20. ANOVA Results – Between-Site Variation in Mean Trap Dates of

Scaphinotus angulatus. Based on online software at
www.physics.csbsju.edu/stats/anova.html.

Source Df Sum Squares Mean Square F Probability

Between 1 6075 6075 0.9068 0.36
Within 14 93787 6699 ---
Total 15 99862 --- ---

Page 22 of 32 -- Cychrine Surface Activity by Kirk

Scaphinotus rugiceps

Scaphinotus rugiceps was trapped at only one site, so between-site comparison is not
possible.

Sex-Related Differences

I compared mean trap dates of males to those of females for the 16 species-site-years for
which sufficient data were generated: two comparisons for S. rugiceps; four for S.
velutinus; and five each for S. angusticollis and S. marginatus. I used external
characteristics to distinguish male Scaphinotus from females: male Scaphinotus have
laterally expanded tarsal segments on the first leg, with dense plantar pubescence;
females have unexpanded tarsal segments on the first leg, with no dense pubescence.
Because I was not able to distinguish sexes in C. tuberculatus based on external
characteristics, and did not take the time to dissect genitalia to determine sex, I made no
male-female comparisons for that species.

If the total number of specimens in a data set exceeded 30, I calculated d (Bailey, 1995);
otherwise I calculated t. With one exception (S. rugiceps at the Fall Creek site in 2005; t
= 5.09, 27 degrees of freedom), the differences in means of males and females was not
significant for any species-site-year at the .05 level.

However, when all 16 species-site-years were considered together, a pattern was

apparent: in 13 of those 16 species-site-years the mean trap dates of females were earlier
than those of males. See Table 21. The differences in means was significant at the .05
level (t = 3.031 with 15 degrees of freedom; table t = 2.131). The mean difference was
22.2.

Table 21. Differences in Mean Trap Dates of Male and Female Scaphinotus. In the
second and third columns are annual Julian dates. In the right column are the numbers of
days by which the mean trap date for females preceded the mean trap date for males. A
negative number indicates that the mean trap date for males preceded the mean trap date
for females.

Species, Location, Difference

and Year Males Females Males - Females
S. angusticollis, 290 279 11
Cape Lookout, 2004
S. angusticollis, 297 292 5
Cape Lookout, 2005
S. angusticollis, 275 256 19

Page 23 of 32 -- Cychrine Surface Activity by Kirk

Sand Lake, 2004
S. angusticollis, 298 283 15
Sand Lake, 2005
S. angusticollis, 210 225 -15
Hamlet, 2003
S. velutinus, Cape 253 239 14
Lookout, 2004
S. velutinus, Cape 237 215 22
Lookout, 2005
S. velutinus, Sand 201 176 25
Lake, 2004
S. velutinus, Sand 263 199 64
Lake, 2005
S. marginatus, Cape 152 137 15
Lookout, 2004
S. marginatus, 175 137 38
Nestucca, 2004
S. marginatus, 125 109 16
Niagara, 2004
S. marginatus, Fall 224 233 -9
Creek, 2004
S. marginatus, 120 106 14
Hamlet, 2003
S. rugiceps, Fall 238 245 -7
Creek, 2004
S. rugiceps, Fall 282 193 89
Creek, 2005

Surface Activity and Precipitation

The five trap-site rain gauges were situated under the forest canopy, and received less
precipitation than if they had been situated in the open. A comparison of the two rain
gauges at the Cape Lookout site shows a nearly linear relationship between open-area
precipitation and under-canopy precipitation (Figure 2). The canopy seems to intercept
roughly one-third of the precipitation, regardless of the intensity of the precipitation.

Figure 2. Comparison of Precipitation Rates in the Open (Roadside) and Under-

Canopy (Trap-Side) at the Cape Lookout Site. The curved line is fitted. The straight
line reflects the linear regression of the trap-side precipitation rates against the roadside
precipitation rates.

Page 24 of 32 -- Cychrine Surface Activity by Kirk

Surface activity and site-specific precipitation measurements are displayed graphically in
Appendix Figures 20 through 31

Because I collected trap-site precipitation data for less than half of the total trapping
period, and because the open-area and under-canopy precipitation rates correlated well, I
explored the relationship between the local, under-canopy precipitation data I collected
and the precipitation data for the same periods from Astoria, Oregon. Astoria was the
closest weather monitoring station with a complete set of data at the time I was analyzing
data. Astoria is approximately 60 miles north of the trap sites. I collected no trapping-
related precipitation data for the Hamlet site.

The regression of Astoria precipitation on local, under-canopy precipitation shows that

the local, under-canopy precipitation is strongly associated with Astoria precipitation (see
Table ). Because of that strong association, and because data are available from Astoria
for the entire trapping period, I chose to use the Astoria precipitation data for part of the
analysis rather than the local precipitation data I collected during less than half of the
trapping period.

Page 25 of 32 -- Cychrine Surface Activity by Kirk

Figure 3. Average Daily Precipitation at Trap Sites and Astoria. Inspection suggests
a strong association between the average daily precipitation during trapping periods at the
trap sites and precipitation records for the same periods at Astoria, located approximately
95 kilometers (60 miles) to the north.

Table 22. Regression Coefficients and Calculated t Values of Astoria Average Daily
Precipitation Against Trap Site Average Daily Precipitation During 26 Trap
Periods. When t is larger than 3.74 (the table value of t with n-2 (n = 26 in this case)
degrees of freedom), the probability of the departure of the regression from zero being
due to chance alone is 0.001 or less.

Calculated t Values
Regression for Regression
Site Coefficient Coefficient
Cape Lookout .971 5.22
Sand Lake Road .965 8.04
Niagara Road .824 8.16
Nestucca Road .800 9.57
Fall Creek Road .874 6.99

Page 26 of 32 -- Cychrine Surface Activity by Kirk

The regression of trap success against Astoria average daily precipitation is summarized
in Table 23. The association between trap success and average daily precipitation is not
statistically significant at the .05 probability level for any of the species. This lack of
statistical significance reflects the high variability of the data.

Table 23. Regression Coefficients and Calculated d Values of Trap Success Against
Astoria Average Daily Precipitation During 52 Trap Periods. When the absolute
value of d is larger than 1.96 (the table value of d), the probability of the departure of the
regression from zero being due to chance alone is 0.05 or less.

Calculated d Values for

Site Regression Coefficient Regression Coefficient
Scaphinotus angulatus .0371 .0242
Scaphinotus angusticollis 5.03 .162
Scaphinotus velutinus -.285 -.0485
Scaphinotus marginatus .343 .0384
Scaphinotus rugiceps -.0900 .0253
Cychrus tuberculatus -1.72 -.246

Although the overall correlation between precipitation and activity is poor, there does
appear to be coincidence of the onset of fall rains and the onset of surface activity in
Scaphinotus angusticollis. See Appendix Figures 21 and 22. There also appears to be
coincidence of peaks in precipitation and peaks in surface activity in Scaphinotus
marginatus. See Appendix Figure 23. Further investigation would be necessary to
determine whether those coincidences reflect some sort of linked relationship between
precipitation and surface activity.

Surface Activity and Temperature

Soil Temperature

Measured soil temperatures are presented in Table 24 and displayed graphically in Figure
4. Soil temperature and surface activity are displayed graphically for the various species
and sites in Appendix Figures 32 through 43. Inspection of those figures reveals no
obvious relationships, with two possible exceptions: (1) Scaphinotus marginatus was
not active when soil temperatures were high in the summer; and (2) Scaphinotus rugiceps
showed a precipitous drop in surface activity as soil temperatures peaked in August, and
a resumption of surface activity as they dropped in late September.

Page 27 of 32 -- Cychrine Surface Activity by Kirk

Table 24. Soil Temperatures. Temperatures were measured just below soil surface, not
more than 1 cm deep, using a 6-inch alcohol thermometer. Readings were recorded to
1/2º C, and are rounded in this table to even number (e.g., 1-1/2 rounded to 2; 2-1/2
rounded to 2). The thermometer was calibrated in ice bath -- 1º on thermometer = 0º C.

Date Cape Lookout Sand Lake Niagara Nestucca Fall Creek

7 Nov 04 nd 10 9 8 9
20 Nov 04 7 6 4 4 4
5 Dec 04 6 6 4 4 4
18 Dec 04 13 13 9 7 9
5 Jan 05 2 2 1 0 0
21 Jan 05 9 10 8 7 9
4 Feb 05 8 9 7 5 7
22 Feb 05 8 8 3 3 3
8 Mar 05 12 12 12 11 12
20 Mar 05 9 9 7 7 6
3 Apr 05 7 7 6 6 5
17 Apr 05 8 7 5 6 5
29 Apr 05 10 10 10 10 9
13 May 05 12 12 12 11 9
29 May 05 12 13 12 12 11
15 Jun 05 11 11 10 8 8
28 Jun 05 13 13 12 13 13
15 Jul 05 10 11 10 11 11
29 Jul 05 11 16 19 17 21
1 Sep 05 13 14 13 11 16
16 Sep 05 12 12 11 11 11
4 Oct 05 9 9 nd 8 8
18 Oct 05 13 14 14 13 13
8 Nov 05 7 7 6 5 5
9 Dec 05 4 4 1 1 1

Page 28 of 32 -- Cychrine Surface Activity by Kirk

Figure 4. Soil Temperatures at Trap Sites

The soil temperature at the time of the first surface activity of each species at each site
was estimated by calculating the mean of the soil temperature measured at the beginning
of the relevant trap period and the soil temperature measured at the end of that period.
Those soil temperatures are presented in Table 25. Analysis of variance using Dr. Robert
Knodt’s MODSTAT software indicates that variances are homogeneous, that the soil
temperatures when Scaphinotus marginatus and S. angulatus first became active on the
surface were significantly lower than those when Scaphinotus angusticollis and S.
velutinus first became active on the surface, and that those when Scaphinotus
angusticollis first became active on the surface were significantly higher than those when
Cychrus tuberculatus first became active on the surface. The t-test scores are presented
in Table 26.

Page 29 of 32 -- Cychrine Surface Activity by Kirk

Table 25. Soil Temperatures (ºC) at Time of First Surface Activity. These are the
means of the soil temperatures measured at the beginning and end of the trap period
during which the species was first trapped.

S. marginatus S. angulatus C. tuberculatus S. velutinus S. angusticollis

5.50 7.5 9 12 12
6.50 5.5 8.5 11 12.5
9 7.5 7.5
8
10

Table 26. t-Test Values from Dr. Robert Knodt’s MODSTAT Software

S. angulatus C. tuberculatus S. velutinus S. angusticollis

S. marginatus 0.325 1.680 3.227 1.930
S. angulatus 2.351 4.472 1.908
C. tuberculatus 3.782 1.596
S. velutinus 0.400

The Duncan Multiple Rang e Test indicates that the soil temperatures when Scaphinotus
velutinus first became active on the surface were significantly higher than those when
Scaphinotus angulatus, S. marginatus, and Cychrus tuberculatus first became active on
the surface, and that the soil temperatures when Scaphinotus angusticollis first became
active on the surface were significantly higher than those when Scaphinotus angulatus
first became active on the surface.

Tukey’s Test and the Scheffe Procedure indicated no significant differences.

These results should be viewed with caution. Because the calculation of mean
temperatures involves only two measurements separated by several weeks, the calculated
mean provides only a very rough approximation of the temperature when a species first
became active on the surface. The availability of only two sets of measurements for
Scaphinotus angulatus introduces another possible source of error. The discrepancies
between the t-test and Duncan’s Test, and the absence of any significance with Tukey’s
Test and the Scheffe Procedure reinforce the need for caution.

Air Temperature

In part because of the uncertainty associated with the soil temperature results, and in part
because use of air temperatures generated more data, I examined the relationship between
the earliest dates of surface activity and air temperatures recorded at Astoria, Clatsop

Page 30 of 32 -- Cychrine Surface Activity by Kirk

County, Oregon, which is located about 60 miles north of the trap transect. Astoria is the
closest source of complete U.S. Weather Service temperature data during 2003 through
2005.

The significance of differences in air temperatures at the onset of surface activity is

summarized in Table 26. The air temperatures at the onset of surface activity of
Scaphinotus marginatus are significantly lower than those of all other species, reflecting
its earlier appearance on the surface. The differences among other species are not
significant.

Table 25. Significance of Differences in Mean Temperatures (ºF) at Onset of

Surface Activity. Absolute value of calculated t value, with table t value at p=.05 in
parentheses. Significant differences shown in boldface. Temperatures are means of the
average daily temperatures reported at Astoria, Clatsop County, Oregon during each
trapping period.

S. angusticollis S. marginatus S. rugiceps S. velutinus C. tuberculatus

S. angulatus 0.99 (2.262) 2.44 (2.262) 0.42 (3.182) 0.55 (2.447) 0.70 (2.179)

S. angusticollis 2.35 (2.145) 0.18 (2.306) 0.60 (2.201) 0.73 (2.110)

S. marginatus 4.31 (2.306) 3.84 (2.201) 4.40 (2.110)

S. rugiceps 0.30 (2.571) 0.28 (2.201)

S. velutinus 0.04 (2.145)

ACKNOWLEDGMENTS

I thank Jenny Kirk and Destiny Kirk for assistance in the field. I thank James R. LaBonte
for encouragement, advice regarding identifications, and a critical review of a draft of
this report. All errors remaining in this report are solely my responsibility.

Page 31 of 32 -- Cychrine Surface Activity by Kirk

 LITERATURE CITED

Bailey, Norman T.J. 1995. Statistical Methods in Biology, Third Edition. Cambridge
University Press, Cambridge, U.K., xiv +255 p.

Gidaspow, Tatiana. 1968. A revision of the ground beetles belonging to Scaphinotus,

subgenus Brennus (Coleopter, Carabidae). Bulletin of the American Museum of Natural
History, 140(3):135-192.

Gidaspow, Tatiana. 1973. Revision of ground beetles of American Genus Cychrus and
four subgenera of Genus Scaphinotus (Coleoptera, Carabidae). Bulletin of the American
Museum of Natural History, 152(2):51-102.

Johnson, Norman E., William H. Lawrence, and Irwin D. Ellis. 1966. Seasonal
occurrence of ground beetles (Coleoptera: Carabidae) in three habitats in southwestern
Washington. Annals of the Entomological Society of America 59(6):1055-1059.

La Bonte, James R. 1998. Terrestrial Riparian Arthropod Investigations in the Big Beaver
Creek Research Natural Area, North Cascades National Park Service Complex, 1995-
1996: Part II, Coleoptera. Technical Report NPS/NRNOCA/NRTR/98-02, United States
Department of Interior - National Park Service - Pacific West Region,
http://www.nps.gov/noca/arthropod2-t3.htm, 138 pages.

Pearsall, Isobel A. 2003. Study to assess the efficacy of ground beetles (Coleoptera:
Carabidae) as ecological indicators in two variable-retention experimental plots: year 2
final report. Forest Investment Account Report No. 6080009, 127 p., photographs,
tables, figures

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