by
Charles W. Baker
James C. Munger
Luana McCauley
Marilyn Olson
George Stevens
IDAHO BUREAU OF LAND MANAGEMENT TECHNICAL BULLETIN NO. 94-1 FEBRUARY 1994
BRUNEAU DUNES TIGER BEETLE INVENTORY
FINAL REPORT
for
Challenge Cost Share Project: D010-P3-0097
Submitted to: Bureau of Land Management
Boise District Office
3948 Development Ave.
Boise, Idaho 83705
January 19, 1994
*n.
Principal Investigator Charles W. Baker, Ph.D.
Department of Biology
Boise State University
Boise, Idaho 83725
James Munger, Ph.D.
Department of Biology
Boise State University
Boise, Idaho 83725
Luana McCauley, Senior Student
Department of Biology
Boise State University
Boise, Idaho 83725
Marilyn Olson, Senior Student
Department of Biology
y ' Boise State University
« Boise, Idaho 83725
George Stephens
Information Manager
Conservation Data Center
Idaho Department of Fish & Game
600 South Walnut
Boise, Idaho 83712
RRIJNEATJ DUNES TIGER BEETLE INVENTORY
TABLE OF CONTENTS
INTRODUCTION 1
OBJECTIVE ONE: DISTRIBUTION OF CICINDELA ARENICOLA IN
OWYHEE COUNTY 2
DISTRIBUTION AT BRUNEAU DUNES STATE PARK 2
A NEW SITE 3
OBJECTIVE TWO: POPULATION SIZE ESTIMATION OF CICINDELA
ARENICOLA FOR SITE C AT BRUNEAU DUNES
STATE PARK 5
PART 1: ADULTS
Introduction 5
Materials and Methods 5
Results 6
Discussion 6
PART 2: LARVAE
Introduction 7
Materials and Methods 8
Results
Density and Total Population Size of Late Instar Larval Burrows 11
Habitat preference of Larvae 12
Discussion 13
MANAGEMENT RECOMMENDATIONS 16
RECOMMENDATIONS FOR GROUP ONE 17
RECOMMENDATIONS FOR GROUP TWO 19
RECOMMENDATIONS FOR GROUP THREE 20
FIGURES 22
MAPS 26
PLATES 30
SUMMARY OF UNITS OF ACCOMPLISHMENT 43
ACKNOWLEGMENTS 44
LITERATURE CITED 45
APPENDIX 46
INTRODUCTION
In the decade of the eighties a tiger beetle now known as The Dunes Tiger Beetle became a species
of concern. The range of this species, Cicindela arenicola Rumpp, is restricted to the dune
systems associated with the Snake River in Idaho. Collectors and naturalists found the numbers
of this species to be greatly reduced in the Bruneau Dunes State Park area. Also, it was no
longer found at some localities where it had been previously collected (Anderson 1 988). In
accordance with the Endangered Species Act, the Dunes Tiger Beetle (Cicindela arenicola
Rumpp) has been designated a Category 2 species.
William F. Barr of the University of Idaho first collected the beetle in 1 963, and Norman L
Rumpp later described it (Rumpp 1 967). Cicindela arenicola is one of sixteen tiger beetles
known to occur in Idaho, and is apparently the only species limited to Idaho (Shook 1 984).
The species' range extends from the general area of the Bruneau Dunes State Park in Owyhee Co.
in the west to the St. Anthony dunes complex in Freemont Co. in the east. The eastern
populations evidence high numbers and appear to be thriving (Anderson 1 989). In contrast,
the western populations occur at much lower numbers and are thought to possibly be on the
verge of extinction (Anderson 1 989).
Cicindela arenicola is reported to have both a spring and fall activity period but is inactive
during the hottest summer months. Adults mate and lay eggs in the spring, and first instar
larvae hatch in late spring or early summer. Both adults and larvae are predators on other
invertebrates (Anderson, 1 988). Adults actively run over open dunes in search of prey and
mates (Plates 1 & 2). Larvae live in vertical burrows where they wait for passing prey.
Larval burrows are more common in the more stable, flat areas where the sand is not actively
drifting (Plates 3 - 5). Larvae require two to four years to complete their development during
which time the burrow is periodically excavated to a larger size and depth (Bauer 1991).
Piles of small balls of excavated material are a great aid in finding these burrows which
otherwise blend very well with the background, especially when the head capsule of the waiting
larva fills the burrow opening (Plate 6).
This study had two major objectives:
1 . to determine the distribution of Cicindela arenicola at this time within Owyhee
County.
2. to estimate the size of the population of £. arenicola at Bruneau Dunes State Park.
OBJECTIVE ONE: DISTRIBUTION OF CICINDELA ARENICOLA IN OWYHEE COUNTY
INTRODUCTION
Previous reports indicated that Cicindela arenicola existed in very low numbers in a very
limited area in Owyhee County. The only known sites were within, or bordering Bruneau Dunes
State Park (Anderson 1 989, 1 992). These previously known sites and other prospective areas
within the park were checked by foot surveys. All terrain vehicles were not used due to the
fragile nature of the habitat and the wary nature of the beetles. Foot surveys were also
conducted in other areas of Owyhee County far removed from the park. Areas outside the park
were selected for survey using soil maps provided by Jim Klott of the Twin Falls Office of the
Bureau of Land Management. Surveys were conducted in areas with sandy soil in hopes that
these areas might provide conditions adequate to support populations of the beetle. As a result of
these surveys the Dunes Tiger beetle is now known to occur at two sites in Owyhee County: the
previously known Bruneau Dunes State Park area and the newly discovered Windmill Site,
located about 1 3 km east of Bruneau Dunes State Park.
DISTRIBUTION AT BRUNEAU DUNES STATE PARK
Surveys conducted within the park established the existence of one large area (Site C below) and
several limited areas wherein smal1 .-populations of the beetle occur. Areas of suitable
habitat are generally located in the rather flat transitional zones situated between the barren
sand dunes and the surrounding desert vegetation. Cicindela arenicola had been seen in some of
these areas in the past, but generally in quite low numbers. Historically, collectors favored the
smaller subpopulations that occurred along the old boundary on the western side of the Park.
These sites were designated populations A and B by Anderson in his 1992 report (See Plates 7-
1 0). Collectors indicate that historically these sites had much higher numbers of adults and
were, therefore, those sites more often frequented for the collection of adults of this species. (G.
A. Shook, personal communication). Site C is situated about 1.5 km southwest of subpopulation
A. It is the largest subpopulation at Bruneau Dunes State Park with about 84,250 sq. meters of
larval habitat (see Map 1).
The smaller subpopulations are principally distributed along the eastern and southern edge of
the major dune system of the Bruneau Dunes State Park. They display a patchy distribution.
The scope of this report does not include measurement of these habitat patches or comprehensive
surveys for the estimation of the numbers of adults and larvae in each area. Their collective
numbers appear lower than those for the single, larger subpopulation at Site C. George
Stephens of the Conservation Data Center conducted the surveys which pinpointed many of these
smaller communities at Bruneau Dunes State Park. Copies of George Stephens' field notes are
included as an appendix of this report. All locations where £. arenicola is known to occur at the
Bruneau Dunes State Park are given on the map (see Map 1).
A NEW SITE
Surveys conducted at locations outside of Bruneau Dunes State Park disclosed the presence of a
previously unknown population of the Dunes Tiger Beetle. This new locality is about 1 3 km
east of the park and has been named the Windmill Site since a windmill is located nearby (see
Plates 1 1-16). The area is accessed from Pot Hole Road south of Hammett in Owyhee County
(see Map 2). Surveys conducted for this area disclosed very limited numbers of both adults and
larvae.
The Windmill Site represents a distinct biological population well isolated from other known
populations in the western portion of this species' range. The inoperative windmill at this site
may have served to concentrate cattle activity in past years. Cattle activity has been shown to
have a potentially negative impact on larval survival of Cicindela arenicola (Bauer 1991).
This beetle population may have been adversely impacted in the past by cattle routinely
traversing the larval breeding areas on their way to water.
Time limitations precluded an extensive inventory of the Windmill Site population this year.
Three walk-through censuses to assess adult numbers were done. No more than 10 beetles were
seen on any one of the surveys. Three adult beetles of £. arenicola were collected as voucher
specimens for this site on May 1 , 1 993. The area in which larval burrows are known to occur
is estimated to be about 500 m long by 50 to 75 m wide. This is not a very big area to sustain a
population. The area where larval burrows occur runs generally north to south in a depression
along the eastern edge of the largest dune complex at this site. Areas of larval habitat have a
quite patchy distribution within this general area. The brief survey conducted on May 1 for
larval burrows indicated that they were at a lower density here than at site C at Bruneau Dunes
State Park. There was evidence of some off-road vehicle use at this site with a few old tire
tracks running through the larval habitat area.
A few specimens of a related species, Cicindela tranquebarica. were also collected at this site.
Larval habitat for £. tranquebarica is found within this dune system where the wind has exposed
an area of saline clay substrate with a high soil moisture level. This was the only other species
of tiger beetle collected on the visits to the Windmill Site this year. Photographs of this site are
provided in the appendix.
OBJECTIVE TWO: POPULATION SIZE ESTIMATION OF CICINDELA ARENICOLA FOR
SITE C AT BRUNEAU DUNES STATE PARK
PART 1 : ADULTS
Introduction
A mark and recapture technique was used to develop a population estimation for adults of £.
arenicola in the eastern portion of its range (Anderson 1 989). The very low numbers of adults
seen in the western populations required the use of a different technique for the estimation of
population size. It was deemed better not to stress these few adults by netting, marking, and
attempting recaptures.
Materials and Methods
The surveys for adults started at a marker (stamped C1/4, S27.T6SR6E) near the south end of
this dune system. Three observers walked north in a line on the open dunes with a distance
between them of about 1 0 m. The observer on the eastern end of the line adjusted to
topographical changes so as to remain about 1 0 m from the edge of the open dune system. The
other observers adjusted their line of march to accommodate for changes in direction taken by
the observer on the eastern end. The length for this line of march was about 850 m with a width
of about 40 m. The area surveyed was 3.4 hectares. Surveys were done in the morning when
surface temperatures became optimum for beetle activity (80-1 1 0 ° F). Sighted adult beetles
were counted only when they were passed or they flew behind the line of walkers so as not to
recount the same beetle.
Results
The results of the three surveys are as follows:
DATE TEMPERATURE
Sand Surface Air
May 12, 1993 110°F 94 F
May 13, 1993 85°F 76°F
May 14, 1993 86'F 76°F
NUMBER OF ADULTS
9
8
11
The total area for each of these surveys was 3.4 hectares. The average number of adults seen
during each of these three surveys was 2.7/ hectare.
Discussion
The number or adults seen per hectare at Bruneau Dunes State Park was much lower than the
number seen by Anderson (1 989) in the eastern portion of the range of this species. We saw an
average of 2.7 beetles/hectare. Anderson reported 7.6 beetles/hectare for the smaller, more
heavily vegetated dunes and an average of 14.26 beetles/hectare for the larger, more open dunes
which represented the better habitat for the Dunes Tiger Beetle. It is of interest to note that his
lower values were reported for those dunes which for vegetation and size are more like the
dunes seen at Bruneau Dunes State Park.
The highest number of adults seen on any walking survey for 1 993 was 1 1 beetles. Such low
numbers could easily contribute to significant errors in population estimation. Although our
measures of density did not vary widely over time, we feel little confidence in the reliability of
our measures. This is because surface conditions on the dunes can fluctuate widely within very
short time periods. The beetles adjust their activities according to the prevailing conditions.
Whenever it is too hot or too cold adults will burrow into the sand. At the time of a survey not
all adults may be active, due to the topographical differences at the various places where they
have burrowed into the sand. In surveys of large populations such variability is less important.
With very small populations, however, the absence of just a few individuals could greatly skew
the results. The very wary nature of the adults also made detection difficult. When the wind
was blowing, movement of vegetation and drifting debris made detection of flying adults
virtually impossible.
The very low number of adults, their very wary nature and cryptic condition, and the rapid
fluctuation in conditions which determine their observability suggested that an alternative
method should be employed to estimate the size of the beetle population at Bruneau Dunes State
Park. Accordingly, a method utilizing the number of late instar larval burrows was developed
for population size estimation. Late instar larval burrows are burrows with a diameter of at
least 4 mm and which presumably contain only third instar larvae.
PART 2: LARVAE
Introduction
Late instar larval burrows should provide a more reliable basis for deriving an estimation of
population size. We decided to use the number of late instar larval burrows ( 4 mm diameter
openings) as the basis for population number assessment for the following reasons:
1 . The burrows are regularly available for counting throughout the early spring activity
period. The presence of adults, by contrast, is much more dependent on weather
conditions. The burrows generally remain open under cool, windy, or cloudy conditions.
Such conditions often cause the adults to take refuge beneath the surface where they are
unavailable for survey purposes.
2. Only the largest class size (4 mm) was used to derive a population size estimation
since the smaller burrows are much more difficult to see and could more easily be
overlooked. Use of only the largest size should reduce counting errors.
3. The 4 mm size burrows contain the largest larvae and these are more likely to
successfully reach adulthood. This 4 mm cohort is, therefore, the best size class for
predicting adult numbers in the next year.
4. The burrows have a fixed position which can be mapped and marked and used for other
study purposes if desired.
Larval burrows are available throughout the seasonal activity period of this beetle. They are
apparently opened in the spring at very nearly > ^ same time that adults appear. Burrows
probably have a permanent location once established but larvae are reported to occasionally
abandon a burrow, move, and quickly excavate a new burrow (Pearson 1988). Some burrows
were still active on June 4, 1 993, when the field work for this study was concluded. No adults
were seen after May 1 9 of this year. An estimation of the population size was done by counting
the number of the late instar larval burrows present in systematically placed quadrats and then
extrapolating for the total area of larval habitat.
Materials and Methods
Larval habitat occurs in a narrow band between the drifting sand of the dunes and the established
desert plant community. This ecotone band is characterized by scattered sandy hummocks with
interspersed flat areas of varying size. Sandy hummocks generally support some type of
vegetation (cheat grass, rice grass, sand-bur, and Russian thistle) which serves to hold the
drifting sand. The intervening flat areas have a covering of small gravel, pebbles, or rocks
which helps to stabilize the underlying sand. These flat areas usually lack plants but may
support a limited number of the same species of plants that occur on the hummocks. When
plants are present in the flat areas, they are smaller and appear to be more stressed than those
on the hummocks.
The length of the larval ecotone habitat is approximately 950 m in a north-south direction.
The width of the band in an east-west direction is about 1 00 m, tapering to 50 m in the last
100 m at each end. The band is shaped in a gentle arc (see Map 1). Initial plans envisioned
transects at every 40 m, but time constraints required that some be done at every 80 m if the
entire area was to be surveyed. Eighteen west-to-east transects were established along the
length of the arc. The length of the transect varied with the distance to which larval habitat
extended out from the dune system. Three transects on the ends were only 50 m long. All others
were 100 m long. Transects A through G are 80 m apart. Transects N9 through S1 are 40 m
apart. The distance between transect G and transect N9 is 54 m, and the distance between
transect S2 and S1 is 57 m. (see Fig. 1). The larval habitat area that was studied is estimated
to occupy 84,250 sq. meters.
Each transect started at the edge of the drifting sand that bordered the west side of the larval
habitat. The transects traversed to the east across the ecotone band to terminate in the more
established plant community that bordered the east side of the larval habitat. Transects started
at a point on the dune that was within a few meters of where the dune stopped and the transition
habitat that supported larval burrows commenced. They ended in an area which was beyond the
preferred transition habitat where the larvae normally occur (see Plate 1 7).
Permanent markers (orange-painted digger bars pounded to within a few inches of the surface)
were placed at the starting and ending points for each transect. Additional digger bars were
placed at every 25 m along each transect. The starting point of each transect was flagged, as
were the end points of most transects. Transects were established using a surveyor's Brunton
compass and tripod on a sighting of N30W/S30E. A meter tape was affixed to the zero, 50 and
1 00 m digger bars as a reference line for positioning of quadrats. Every other square meter of
habitat along the north side of the transect was inventoried using a quadrat that was 1 m^- A
10
coin toss determined whether odd or even numbered quadrats on a given transect were
inventoried.
Data was taken for each quadrat by one observer on the south side of the quadrat and another
observer on the north side. Estimations of each variable (see below) were made by each
observer; when different values were obtained a reevaluation was done to the mutual agreement
of the observers (see Plate 1 8).
The information obtained for each quadrat surveyed is as follows:
1 . Number, size, and placement of any larval burrows.
The diameter of the opening of each larval burrow was measured. There were
three classes for the diameter of the larval burrows: 2, 3, and 4 mm. The
particular decimeter square of the quadrat in which each burrow was located
was also recorded.
2. Percent vegetation cover.
Vegetation cover was estimated to the nearest 5%.
3. Nature of the composition of the surface layer.
Surface coverage with sand, gravel, pebble or rock was determined. Sand was
material of less than 1 mm diameter, gravel was from 1 to 3 mm diameter,
pebble was 3 to 13 mm diameter, and rock was greater than 1 3 mm diameter.
Percent coverage within each quadrat was estimated for each of these four
categories.
4. Topography.
The surface was rated as level if it was flat and not inclined, as even if it was
flat but inclined, or uneven if the surface was irregular due to the presence of
hummocks.
11
Great care was taken to minimize the impact of this survey upon the habitat under investigation.
Bauer (1 991 ) had previously demonstrated that cattle walking on burrows could collapse the
burrow tubes of the larvae of this beetle such that they could not recover. Early in this study
experimentation was done with special footwear (sandshoes) that more widely distributed one's
weight, but the footwear proved too cumbersome. A simple compaction test was also conducted
with normal footwear. One person deliberately stepped twice on the opening of two burrows of
the 4 mm diameter category. Although both burrows were successfully reopened by the next
day, it is likely that repeated trampling would reduce survival. A larger sample size was not
used due to the sensitivity of this species. The amount of compression and general disturbance
caused by someone walking on larval burrows could be expected to be greater if the sand was
much drier or if the sand was of a much finer nature. It is also likely that smaller diameter
burrows, especially the 2 mm size, would be less likely to successfully recover from such
disturbance. The period of larval hatching and establishment which occurs in late May and early
June would be a time when these very small larvae with rather shallow burrows would be most
sensitive to surface disturbance.
As the work progressed it became quite evident that burrows of the beetle larvae occurred
nearly exclusively in the flatter, more open areas. We could generally avoid walking on burrow
openings by keeping to the elevated hummocks. When forced to traverse the areas of larval
habitat we went slowly and stepped only where no burrow openings were seen. Anyone following
walked in the footprints of the lead person (see Plates 1 9 & 20).
Results
Density and Total Population Size of Late Instar Larval Burrows
Twelve late-instar burrows were seen in the 825 quadrats, each 1 m2, that were carefully
inventoried. From these data we can estimate first, that the density of burrows is 0.01 45455
burrows/m2 (+/- 95% CI) and second, that the total number of late-instar larval burrows is
12
1,225.5 for Site C, with a 95% confidence interval of 521.5 to 1929.5 (Scheaffer, et al.,
1979). The coefficient of dispersion (variance to mean ratio) for the number of larvae per
quadrat equals 0.987, which gives no evidence of non-random distribution (clumped or
uniform) of larvae, at least at the scale of 1 m^ quadrats.
The design of the larval study was such that all larval habitat was supposed to occur within the
transect grid. During the latter part of the project several small outlying areas of larval
habitat were found just east of the study area for site C. The collective size of these outlying
communities was not deemed large enough to materially affect the study. Our best estimation is
that the total square meter area of all of the other larval habitat at the Bruneau Dunes State
Park including these outlying areas near site C does not exceed the total square meter area
surveyed this year for site C. The total late-instar larval population for Cicindela arenicola at
Bruneau Dunes State Park probably does not exceed 2,500 specimens, with a 95% confidence
interval of 1,043 to 3,859.
Habitat Preference of Larvae
Larval burrows of all ages occurred in only 23 of the 825 quadrats surveyed. For these 23
quadrats there were only 1 2 burrows of the 4 mm diameter size. Larvae were most likely to
occur in quadrats wherein the mix of surface materials was 30% sand, 40% gravel and 30%
pebbles (Figures 2-4).
By using the frequency distributions of the quadrats without larvae as a measure of the available
habitat, we can make several conclusions about the habitat preferences of larvae. First, larvae
occur more than expected in areas of 30-40% sand, and substantially less than expected where
coverage with sand was greater than 50% (Fig. 2). Larvae also were more common than
expected in quadrats with gravel comprising 30-50% of the surface, and were less common
than expected in areas where gravel was less than 10% of the surface (Fig. 3) . Larvae were
13
more common than expected in areas with a 20-40% pebble cover, and less common than
expected where pebble cover was 10% or less (Fig. 4).
Larvae occurred more commonly than expected in areas with little vegetation cover. Over 90%
of the burrows were in areas where the vegetation cover did not exceed 30% and nearly 50% of
the larvae were found in areas with zero to 10% vegetation cover (Fig. 5).
Approximately 60% of the larval burrows occurred in areas categorized as uneven while 40%
of the burrows occurred in the areas designated as even or level. However, since the uneven
habitat is far more common (4.8 times as common), the larvae clearly prefer or survive better
in areas that are even or level (Fig. 6).
Examination of how habitat affinities differ among age classes (Figures 7 and 8) should yield
some insight into the mechanism that causes the preferences thus far discussed. If habitat
preference is expressed by females laying eggs, then the three age classes should have similar
frequency distributions. If females lay indiscriminately and habitat preference is caused by
higher survival in some habitats than in others, then the age classes should have different
frequency distributions, with 2 mm larvae distributed similar to plots with no larvae and 4 mm
larvae showing a distinct preference. The actual result is closer to the latter, although low
sample sizes make definitive determination impossible. Two mm and four mm larvae both
appeared to have higher than expected occurrences at low percent of vegetation, indicating that a
preference for non-vegetated areas is exhibited at the egg-laying stage.
Discussion
The density of burrows for the Dunes Tiger Beetle at Site C was quite low compared to that seen
for several other species of tiger beetles in this part of Idaho. Larval densities for the
populations of this beetle in eastern Idaho have not been reported. It is quite possible that they
14
are higher than those seen for western populations since the adults are reported to have much
higher densities in the eastern populations as discussed previously.
The reliability of estimates made during a single year is questionable. First, the spring of
1 992 was exceptionally wet and cold, and a phenological delay of two to three weeks was common
for most events. The effect of this on adult beetle activity and density is difficult to estimate.
Second, Cicindela arenicola requires two to four years to reach the adult stage (Bauer 1991).
Accordingly, populations of adults may naturally have high and low years which are the result of
good or bad habitat conditions over the one to three years preceding a count. The number of
larval burrows could be affected in a similar fashion. To accurately or meaningfully inventory
the population and assess population trends would require five or more years of annual
monitoring for a species such as C. arenicola.
Late in the study (May 1 8 to June 4) after most of the survey work had been completed, high
numbers of 2 mm diameter burrows appeared at many places within the area of Site C. The high
number of these burrows (which are made by hatching larvae) indicated that a great amount of
ovipositional activity had occurred even though the number of adults observed on any day was
quite low. This would have occurred if peak adult activity occurred this year at times when no
surveys were conducted. Another possible explanation is that multiple year diapause is known
for certain stages of some insects so there is also the possibility that eggs of this species may lie
inactive in the sand for more than one year before optimal conditions stimulate hatching. This
would most logically occur in areas where the surface was stable from year to year such as the
flat areas with gravel and pebbles. These areas would provide more protection for eggs than the
open, changing dune habitat.
Anderson (1 988) noted that the widespread occurrence of new larval burrows at the time of
hatching reflected an apparent non-selective ovipositional behavior for females of the eastern
15
populations. By contrast, the new 2 mm diameter burrows at Bruneau Dunes State Park
appeared almost exclusively within the flat areas, very few being found in the very sandy areas
(see Plate 21 ). The analysis of the habitat data provides additional support for selectivity of
habitat in choosing oviposition sites since 2 mm larvae tended to occur more than expected in
areas with moderate sand and low vegetation cover.
Anderson (1 989) indicated that this beetle did not occur on many of the dunes in Power Co. that
were overgrown with vegetation. Such dunes were deemed by him to be "unsuitable for£.
arenicola." The beetle apparently needs naked dunes in order to exist. We concur with him on
this point and suggest that the low numbers of beetles at Bruneau Dunes State Park may
primarily be due to vegetational encroachment upon the critical habitat for this species. Two
exotic species of plants, Russian thistle and cheat grass, are quite common in much of the
habitat where the larvae are found.
Overgrowth of these areas by weedy species could result in lower survival of larvae since they
have evolved to live in open areas. The presence of vegetation would hinder burrow maintenance
and could make the larvae more vulnerable to predators, parasites, and parasitoids. Vegetation
could prevent the larvae from being able to throw out the sand balls, and vegetation situated
directly above the burrow opening could complicate prey recognition by the tiger beetle larvae.
This could interfer with effective capture of the prey. Shading could also adversely impact
laraval success (see Plate 22).
Increased vegetation would also provide more cover and protection for lizards which can be a
significant predator of adult tiger beetles (Pearson 1 988 and Anderson 1 990). It is quite
possible that the beetles seek the open dune areas for foraging and mating so as to avoid
predation by lizards. Leopard lizards were often spotted taking refuge in the vegetation of the
sand hummocks. The female beetles are believed to enter the flats among the hummocks in order
16
to find ovipositional sites. It is of interest that this species was found most abundantly this year
on these ovipositonal sites at the time of the first visitation. This was on April 20 with an air
temperature of only 64 ° F. The lizards were not active under these conditions. We suggest that
the eariy activity of this beetle may be an adaptation to avoid the increased risk of predation by
lizards that comes with the warmer temperatures later in the spring.
It is the opinion of the writer that the most critical factor in the survival of £. arenicola at this
time is the availability of adequate and undisturbed non-vegetated habitat that will support
larval development for a two to four year period of time. This habitat needs to be in the close
proximity of open dunes where the adults can forage and mate.
MANAGEMENT RECOMMENDATIONS
The several communities of the Dunes Tiger Beetle which occur in Owyhee Co. present a range
of challenges if their continued viability is to be ensured. These populations and subpopulations
can be formed into three groups for purposes of future monitoring and management.
Group one is a collection of smaller and rather scattered subpopulations within Bruneau Dunes
State Park. These disjunct subpopulations occur in a general arc to the east, south and west of
the main dunes complex within the park. Populations A and B in the reports written by Dr.
Robert Anderson are included in this group.
Group two is the extensive subpopulation that is located along the east side of the low dunes
complex located above the bluffs at a higher elevation in the southwest corner of Bruneau Dunes
State Park. This area is also referred to as Site C or Population C in earlier reports.
Group three is the newly discovered Windmill Site. The beetles at the Windmill Site certainly
qualify as a true biological population. The very isolated nature and very small size of this site
17
probably precludes regular genetic exchanges from occurring between this population and other
populations.
Recommendations for Group One
It is recommended that present management practices continue for these scattered
communities. It appears that the habitat available for successful larval reproduction is quite
scattered and quite limited throughout the range of this group. It is very difficult to assess the
impact of introduced species of plants on the vegetation pattern for these scattered areas; it is
unknown if these species are increasing, decreasing, or stable. Such species as Russian olive
trees and Russian thistle have certainly impacted the area and have produced changes in the
landscape which may have caused the perceived decline in numbers of this species.
Vegetational overgrowth is probably deleterious to larval success. The dunes tiger beetle
appears to need rather flat, open habitat for larval reproduction and is apparently adept at
locating such areas as they form. This permits the species to colonize new areas to replace those
that are no longer adequate due to changes in vegetation and dune topography. It is very probable
that historically there have been short-term and long-term fluctuations in the population
numbers of this species. Accordingly, it is virtually impossible to determine in one year
whether or not current numbers of this species are really depressed due to man's impact or are
at a low in a cycle that results from climatic fluctuations and natural biological factors.
It is unknown at this time if cattle or humans have adversely impacted the newly hatched larvae
of this species during the hatching period when they are establishing their first larval burrow.
This critical period occurred during late May and early June this year. It is generally quite hot
by this time in the season and the elevated temperatures tend to reduce park visitations. The
more isolated and less attractive areas which serve as breeding areas are rather infrequently
used by visitors at this time. It is recommended that the use of these areas during this period,
18
especially by large groups, be discouraged. Should burrows be trampled by humans during
other times of the year there is the possibility that at least the more mature larvae would be
able to reopen the burrows. The impact of repeated trampling by humans, however, is
unknown.
In the event that serious negative impacts brought about the demise of one or more of these
scattered subpopulations there is a very high probability that recruitment from other nearby
subpopulations would bring about recolonization. It is even possible that adults move about
among several of these nearer areas. Should all of the subpopulations of this group gradually
decline and fail, re-establishment would be possible from the larger subpopulation at Site C
which is situated on higher ground southwest of the general park area.
It is recommended that the known larval habitat areas for group one be surveyed for the
presence of mature larval burrows at least every two years so as to document the occupation of
these sites by the dunes tiger beetle. The number of adult beetles seen on each of these surveys
should also be recorded. Such survey work would complement that done this year and would
provide data for monitoring the general trend in numbers for these scattered communities.
These surveys should be conducted when weather conditions are proper for adult activity.
It is further recommended that the surveys be done when the sand lily (Leucocrinum montanum
Nutt.) begins to flower (Plate 23). The initiation of activity by the dunes tiger beetle adults
appears to coincide closely with the initiation of flowering by the sand lily. The monitoring of
this plant would appear to provide a good guage for when the beetle commences surface
activities. Calendar dates are less reliable since seasonal variation due to weather conditions
can significantly advance or delay biological activity patterns. Adult beetles were seen on dunes
at Site A on March 22 this year. No flowers were open on the sand lily although buds were
evident.
19
Recommendations for Group Two
Group two is synonymous with population C of Anderson's report (1 992). It is the largest
subpopulation known for the Dunes Tiger Beetle in the western reach of its range. It was first
discovered on April 28, 1 992 at the time of the workshop by Anderson on this species. Plans
call for the inclusion of this area within the boundary of the Bruneau Dunes State Park. The
construction of a fence along the western and southern boundaries by the end of the 1993
calendar year will provide initial protection for this site. This subpopulation will then be
included within the area protected by Bruneau Dunes State Park. The fence should efectively
exclude cattle and off- road cyclists.
This dune system did not appear to be highly impacted by human activities during the spring
months of this year. During the course of the survey work done from April to June of 1993 no
humans were seen to visit this area. A few old wheel tracks from prior years were still visible.
It is quite unlikely that hikers would utilize these more remote dunes. The low and rolling
nature of these dunes offers little in the way of a recreational challenge for hikers. These dunes
are not visible from the more highly developed recreational areas of Bruneau Dunes State Park
situated at a lower elevation to the east.
Subpopulation C is well located for continued inventory and assessment studies since it is not
generally impacted by human activity. Such studies would provide data so as to establish
population trends and elucidate species' dynamics. This area already has an established,
extensive transect system for the purpose of estimating the mature larval population. Annual
counts of larval burrows should be done for at least the next five years so as to establish the
population trend. Counts should also be done for adult beetles at the same time even though they
are less reliable for reasons discussed above.
20
Subpopulation C may well represent a "source site" as discussed by Hubbell and Foster (1 986).
All other sites in the Bruneau Dunes complex may, at least in the short term, represent "sink
sites". The larger, more successful subpopulation C is well-positioned to produce colonizers
for the patchy larval habitat found downwind. Subpopulation C may be the stable reproductive
base that persists while the other areas may come and go.
All the subpopulations within the Bruneau Dunes State Park should be considered as one
biological population since they are probably close enough to permit regular genetic exchange,
considering the known flight capabilities and habits of other species of tiger beetles. Adults of
normally diurnal Cicindela marutha are known to oviposit at night on dunes up to 1 km away
from their diurnal pond edge habitat, and specimens of Cicindela trifasciata have been collected
at lights on offshore platforms 1 60 km from the nearest land (Pearson 1 988).
Recommendations for Group Three
The Windmill Site should be carefully inventoried in 1 994 in view of its small size and
unprotected nature. The area needs to be carefully walked and mapped so as to delineate all the
currently active larval habitat areas. Surveys for both adults and mature larvae should be
conducted to establish a baseline for the numbers of both categories occurring in this
population. At a minimum, three walk-through censuses for adults should be done during the
flowering period of the sand lily (Leucocrinum montanum Nutt.). A transect study should be
performed so as to estimate the number of mature larvae for this small population. A count of
first instar burrows should be done at the time that the first instar larvae hatch and establish
their first burrows. Such counts may prove to be valuable in determining the collective
ovipositional activity in any year.
21
The density of the first instar larval burrows may be a very good indicator for the viability of
populations of this beetle wherever it occurs. Observations at Site C this year indicated a great
overproduction of first instar larvae relative to the number of later instar larvae that survive.
There is the possibility, however, that eggs laid in one season may not hatch until the next year
if hatching conditions are not proper for this species. Thus, there may be an accumulation of
eggs for a few years with extensive hatchings in years of very heavy spring precipitation.
Careful studies need be performed to resolve questions such as this.
If studies at the Windmill Site establish that these dunes are subject to human disturbance such
as camping, offroad vehicles, or all terrain vehicles, then an exclosure fence should be
established to protect the very limited area of habitat that supports larval development.
This population is of considerable biological interest in that it is so small and so isolated. Its
small size allows for ready inventory of the total population at relatively little cost. It is not
recommended, however, that more extensive ecological studies be done at this site, at least in
the near future. Disruption associated with such studies could overtax this site. Any studies
involving manipulation of plants, predators, or parasitoids would more appropriately be
assigned to the larger Site C area of the Bruneau Dunes State Park.
22
Transects
B
nWNWN
WNNNNN
sNNNNNN
ANWW
A\S\V\
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ASWW
sNNNNNN
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i. W WW
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* W W W
tNNW
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s WW w
nWNNW
v W WW
..NNNNNN
1 (18)
Figure 1
Schematic of area surveyed for larval
burrows of Cicindela arenicola Rumpp
at Site C. Transects A, B, and S2 are
each 50 m long. All other transects
are 100 m long. The distance between
transects is to scale in this figure.
WNN
WW
WW
WW
WW
WW
\NNN
^\S\
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LlAA
WW
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; ■■.;>;.
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NNNNNNNNN
NNNNNNNNN
NNNNNNNNN
NNNNNNNNN
NNNNNNNNN
NNNNNNNNN
NNNNNNNNN
Bold numbers
Number in
parenthesis
number of 4 mm diameter
burrows seen in that
transect.
= number of the quadrat in
that transect where the
burrow was found.
WNNNNN
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WNNNNN
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1 (21)
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1 (37)
2(66,30)
1 (61)
2(12,26)
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1C
1
5)
(96,98)
(4)
□ Ab..nl (N .402)
S3 Prri.nl IN . 23)
(C > IQ.9S2, P ■
w
■ '■
n
■■y/A
4 0 50 60 70
Percent Sand
80
I
90
100
23
o :o
C
a 20 ■
%
G Ab<*nt (N a 30?)
3 J'lwnpl.ai
(G > 14. SJ, p > : cu:i
^
i
Figure 3
10 2 0 3 0 4 0
Percent Gravel
50 60
Figure 4
10 20 30
Percent Pebbles
40
24
CO
CL
C
0)
o
CL-
io -
i
!v:
D Absent (N z. 802)
E3 Presenl (N r 23)
_i
II.
i=i
0-10 11-20 21-30 31-40 41-50 S1-S0 61-70 71-80 81-30 91-100
Figure 5 Percent Vegetation
Q.
Q.
Figure 6
D Ablanl (N ■ 802)
0 Pr«s«n( (N > 23)
(G s 11.47, P a .003)
r
Even
Level
Topography
Uneven
25
Q.
C
o
a.
CO
GL
c
CD
O
•—
a
Q.
40-
D Absent
O 2 mm larvae
0 3 mm larvae
£ mm larvae
Percent Cover with Sand
Figure 7
50 i
□ Absent
□ 2 mm larvae
E2 3 mm larvae
E3 4 mm larvae
H—EL
0-9 10-19 20-29 30-39 40-49 50-59 S0-S9 70-79 80-3290-100
Percent Cover with Vegetation
Figure 8
Map 1 - Key to Localities at Bruneau Dunes State Park
1. Subpopulation C This is the largest habitat area for Cicindela arenicola for the area of
this map. It is about 950 meters long and 100 meters wide, bounded on the west by
an open, low dunes system and on the east by a desert shrub community. Charles Baker, Luana
McCauley, and Marilyn Olson. March to June, 1993. Fewer than 2,500 larval burrows are
estimated to occur at this site.
2. A new locality. Twenty larval burrows in small flats bordering low scattered dunes.
Charles Baker and Luana McCauley, May 26, 1993. Six burrows seen here by George
Stephens on June 8, 1993. Fewer than 100 larval burrows estimated to occur at this site.
3. A new locality. Ten larval burrows in a small transition area on the west flank of the big dune
at the edge of the rim overlooking the Park. George Stephens, June 8, 1 993. Fewer than 1 00
larval burrows estimated to occur at this site.
4. Subpopulation A. This is the second largest habitat area for Cicindela arenicola in the vicinity of
Bruneau Dunes State Park. The larval habitat patches are scattered over an area about 200 to
400 meters long and 50 to 100 meters wide. The larval habitat is located along the east side of
a small line of low, open dunes. Charles Baker, Luana McCauley, Marilyn Olson. March to
May, 1993. Fewer than 500 larval burrows estimated to occur at this site.
5. Subpopulation B. No adults or larvae were seen here in 1993. No larval burrows were seen
here in 1992. Mating adults were seen here by Charles Baker in 1992. Cicindela arenicola
does not appear to currently occur at this site.
6. A new locality. One mating pair of beetles on an open sand slope above a small flat about 1 0
meters square wherein there were at least ten larval burrows. It was initially designated as
Population D in notebooks and student reports for 1993. Charles Baker, Luana McCauley,
Marilyn Olson, and George Stephens, May 1, 1993. Fewer than 200 larval burrows estimated
to occur at this site.
7. A new locality. Two adults, no larval burrows. Adults were right at the interface of the bare
sand and the gravel flat. George Stephens, May 19, 1993. Probably no burrows at this site.
8. A new locality. Five larval burrows, no adults. Small area of excellent larval habitat. George
Stephens, May 19, 1993. Fewer than 100 larval burrows estimated to occur at this site.
9. A new locality. Three or four adult beetles on sand slope near small area of what appeared to be
excellent larval habitat, no larval burrows seen. George Stephens, May 19, 1993. Fewer than
50 larval burrows estimated to occur at this site.
10. A new locality. One adult, two larval burrows in an area with small pockets of marginal to
excellent larval habitat. George Stephens, May 19, 1993. Fewer than 100 larval burrows
estimated to occur at this site.
27
Area of Bruneau Dunes State Park
Bruneau Dunes Quadrangle
&
Hot Springs Quadrangle
Idaho
Map 2 - Location of Windmill Site Population
Windmill Site. This population is currently known only from an area that is about 500 meters
long and 50 to 100 meters wide. It runs generally north to south in a depression along the
eastern edge of the largest dune complex in this system. The larval habitat has a very patchy
distribution and varies from marginal to excellent in quality. It is estimated that there are
fewer than 500 late instar larval burrows at this site.
To access this site take State Route 78 south from Hammet to Pot Hole Road which borders the
west edge of this map. Turn off Pot Hole Road onto the road marked in red on this map to
directly access the dune system at the Windmill Site. Agricultural development has changed
some of the roads in this area since this map was made.
29
"T v. ( C
_ Area of Windmill Site
Indian Cove Quadrangle
&
Pence Butte Quadrangle
Idaho
30
PLATE 1
Adult beetle of Cicindela arenicola Rumpp on a sand flat area amongst sand
hummocks at Site C, April 20, 1 993, 64° F, Bruneau Dunes State Park.
*&?mm
Plate 2
Mating pair of adult Cicindela arenicola Rumpp on open dune habitat at
Site D, May 1 , 1 993, Bruneau Dunes State Park.
31
Plate 3
Opening of larval burrow (4mm size) of Cicindela arenicola Rumpp at
Site A, April 1 3, 1 993, Bruneau Dunes State Park. Note mix of sand,
gravel, and pebble surface cover.
Plate 4
Opening of larval burrow (4 mm size) of Cicindela arenicola Rumpp at
Site D, May 1, 1 993, Bruneau Dunes State Park. Substrate here is of a
different color than that at Site A.
32
Plate 5
Opening of larval burrow (4 mm size) of Cicindela arenicola Rumpp at
Site C, April 20, 1993, Bruneau Dunes State Park. Substrate with
pebbles of mixed colors.
p*
Plate 6
Opening of larval burrow (4 mm size) of Cicindela arenicola Rumpp at
Site C, April 20, 1 993, Bruneau Dunes State Park. Note pile of sand balls
thrown out when larva cleans out or enlarges the burrow.
33
Plate 7
Sites A and B at Bruneau Dunes State Park. Picture taken from edge of rim
to southwest that overlooks these sites. Site A indicated by yellow dot.
Plate 8 continues to the right. May 26, 1 993.
_,S9*J^-~"«r->
&•- •wranWflllfcS*- i^^-i-:
?% ***.>•»— -*ss*?aw.-? ^^^^^
^^^^
.
"
- - dJ —
-'■'"
-
■ - - • <*..' . "
v Hp
*■-»*.
J
- - :* >»V>-4--y.*r 4&r»^jfci
•^^"
"". '«
-v '3EJi.tTs^Si
v\T*
*i» -'
"•'--.
■«_
f <-V" ' ; ~ - -:-- '
-
»
~-
*"iC
Plate 8
Sites A and B at Bruneau Dunes State Park. Picture taken from edge of rim
to southwest that overlooks these sites. Site B indicated by yellow dot.
Some of site B can be seen in Plate 7. Trees in center are main campground
May 26, 1993.
34
„<•«
X
*afc
• ;
■T- £"•" ' "!?Sk^";r^^ ,iS
.TL~»»..
Plate 9
Site A at Bruneau Dunes State Park. April 1 3, 1 993. Flag marks
burrow in a flat area amongst sand hummocks. The clump of Russian
olive trees at the upper left corner can also be seen in Plate 7. These
trees were not in leaf on April 1 3, but are seen in full leaf on May
26. In Plate 7 they are seen as the isolated clump of trees at the east
(right) end of the light colored strip of sand that is Site A as marked
by the yellow dot. Most of the larval burrows at this site occur at the
eastern end of the strip near the clump of Russian olive trees. This
site has the second largest area of larval habitat known at this time
for Cicindela arenicola at Bruneau Dunes State Park.
35
Plate 10
Site B at Bruneau Dunes State Park. April 1 3, 1 993. No adults or larvae
were seen here in 1 993. Wind activity appears to be actively eroding
features at this site. The "E.T." column seen in this picture crumbled
in this year. Very little vegetation is seen here and there are few, if any,
stable areas where larvae could live. The pictures seen in Plates 7 and 8
were taken from the edge of the rim in the center background of this picture.
Site C is located about 1 km beyond the rim edge.
36
Plate 1 1
Windmill Site. March 6, 1992. The line of dunes mark the area where
adult beetles were collected later in the year. The Windmill is situated to
the west of this dune line, behind the photographer.
Plate 1 2
Windmill Site. March 6, 1 993. This site has many areas of dunes that
support little vegetation. The sand is of a generally darker brown color
as compared to the color of the sand at Bruneau Dunes State Park.
37
Plate 1 3
Windmill Site. September 17, 1993. The area in which larvae of
Cicindela arenicola Rumpp occur at this site borders, and is
interspersed in, the area of light-colored rice grass seen in the center
of the photograph. Seen here is the north end of the band of rice grass
habitat that continues through Plates 14, 15, and 1 6.
Plate 14
Windmill Site. September 17, 1993. The rice grass habitat that
starts in Plate 1 3 continues in this photograph. Larval burrows of
Cicindela arenicola Rumpp are more abundant in the northern half
of this band of rice grass habitat.
38
Plate 1 5
Windmill Site. September 1 7, 1993. Seen here is the central reach
of the rice grass habitat that borders the area of larval habitat of
Cicindela arenicola Rumpp at this site. This photograph overlaps the
one seen in Plate 1 6.
Plate 1 6
Windmill Site. September 17, 1993. The south end of the band of
rice grass habitat can be seen to spread over the slope of the hill in
the center of this photograph. Very few burrows of Cicindela arenicola
were seen in this area. The exposed saline soil at the lower right
supports a few burrows of Cicindela tranquebarica.
39
Plate 1 7
Site C at Bruneau Dunes State Park. May 28, 1 993. The open,
elevated dune habitat is seen in the foreground and ends near where
the survey tripod is located. Vegetation was abundant this year
due to the much above normal spring precipitation. The foreground
plants are almost exclusively sand bur and Russian thistle.
40
Site C at Bruneau Dunes State Park. May 1 3, 1 993. Transects
started in the very unstable drift sand at the edge of the dune and
preceded out to the more densely vegetated area seen beyond the open
area traversed by the tape. This band of more open habitat in the
middle of the photograph is the area where most larval burrows occur.
t.
Plate 1 9
Site C at Bruneau Dunes State Park. May 1 2, 1 993. Care was taken to
avoid trampling larval burrows in the stable flat areas amongst the
sand hummocks. This larval habitat ceased rather abruptly at the base
of the open dunes seen in the background.
Plate 20
Site C at Bruneau Dunes State Park. May 1 2, 1 993. A flat area near
the north end of the dune system with a very good density of late instar
larval burrows (flags). Few burrows were found in the area beyond
the scratch line in this photo where cheat grass was abundant.
Plate 21
Site C at Bruneau Dunes State Park. June 4, 1 993. The circle of pink
flags encloses an area where more than 60 newly established larval
burrows (2 mm diameter) were seen. Thirty of these burrows were
flagged with green markers for future reference. New burrows were
not seen on the elevated, more vegetated sandy hummocks.
42
' ••-*•-*. r- £&>-~4. •-— i.C?£*-s-
Plate 22
Site C at Bruneau Dunes State Park. May 1 8, 1 993. The edge of the
dunes seen here is heavily overgrown by plants. Larvae may occur
here but their detection is virtually impossible. It is also very
improbable that female beetles would oviposit in such an area.
*8E*!
Plate 23
Near Site A at Bruneau Dunes State Park. April 1 3, 1 993. Seen
here is the sand lily (Leucocrinum montanum Nutt.) at peak bloom.
Adult beetles of Cicindela arenicola Rumpp were seen at site A on
March 22 this year, shortly before the start of flowering by the
sand lily.
43
SUMMARY OF UNITS OF ACCOMPLISHMENT
1 . One new population of Cicindela arenicola was found in Owyhee County and the locality
designated as the Windmill Site.
2. Seven "new" areas of beetle occurrence (either larvae, adults, or both) were
documented for Bruneau Dunes State Park.
3. A transect/quadrat study of Site C at Bruneau Dunes State Park indicated a late instar
larval population of 1 225.5 with an estimated late instar larval population of 1 ,000 to
4,000 for the entire Bruneau Dunes State Park area.
4. Surveys for adult beetles were done at both the Windmill Site and Bruneau Dunes State
Park. No more than 1 1 adults were seen on any survey in 1 993.
5. Surveys of late instar larvae were done at Site C. They showed that the burrows have a
non-random distribution. They are found almost exclusively in rather flat areas with a
mixed surface of sand, gravel and pebbles.
6. Distribution maps for all sightings for 1 993 are included in this report.
7. Management recommendations are given for all known areas where this beetle c jrs in
Owyhee County.
8. Photographs are included in Luana McCauley's report and in the Final
Report for 1993.
9. Copies of the reports done for the Independent Study Projects by Luana McCauley and
Marilyn Olson (Boise State University senior students) are provided with the Final
Report.
44
Acknowledgments
A sincere thank you is given to the many people who provided assistance during the course
of this study:
Jim Klott, Twin Falls District Office of the Bureau of Land Management for assistance
with maps and logistic support.
Ann DeBolt, Boise District Office of the Bureau of Land Management for consultation and
logistic support.
Wes Whitworth, Director of Bruneau Dunes State Park, for resource information and
assistance in locating certain sites.
Dr. Robert Anderson, Biology Department, Idaho State University for a copy of The Idaho
Dunes Tiger Beetle video and technical advice.
Mike Pellant, Christine Hall, and Marlene Ano, Boise District Office of the Bureau of
Land Management for digger bars to mark the transects.
Bill Clark and Gary Shook of the Orma J. Smith Museum of Natural History, Albertson's
College for the loan of reference specimens of several species of tiger beetles.
McDonald, Instrumentation Engineer, Boise State University, for fabrication of
field equipment.
Dr. Marcia Wicklow-Howard, Biology Department, Boise State University, for loan of
field equipment.
Luana McCauley, Senior Student, Boise State University, for photographs.
Ginger Beall, Ann Herndon, and Robert Higdem, Boise State University, student
volunteers.
Tamsen Baker, Meggan Baker, and Liz Baker for technical help.
A special thanks goes to all those listed on the title page who assisted with the field work, and
particularly to Dr. James C. Munger for analysis of the data.
45
Literature Cited
Anderson, R. C. 1 988. The dunes tiger beetle. Final report for BLM Contract ID-030-CT8-
001. Bureau of Land Management, Idaho Falls District.
1 989. The dunes tiger beetle. Final Report for BLM Contract ID-030-CT8-005.
Bureau of Land Management, Idaho Falls District.
1 990. The Idaho sand dunes tiger beetle. Video. Idaho State University Audio Visual
Center, Pocatello, Idaho.
1 992. Observations on the occurrence of the Idaho sand dunes tiger beetle, Cicindela
arenicola Rumpp, in and around Bruneau Dunes State Park on April 28, 1992. A
workshop report submitted to Bureau of Land Management, Boise District.
Bauer, K. L 1991. Observations on the developmental biology of Cicindela arenicola Rumpp
(Coleoptera: Cicindelidae). Great Basin Naturalist 51:226-235.
Hubbell, S. and R. Foster. 1 986. Commonness and rarity in a neotropical forest:
implications for tropical tree conservation, in Michael E. Soule, ed. Conservation
biology: the science of scarcity and diversity. Sunderland. Sinauer.
Pearson, D. L. 1 988. Biology of tiger beetles. Annual Review of Entomology 33: 1 23-1 47.
Rumpp, N. L 1967. A new species of Cicindela from Idaho (Coleoptera: Cicindelidae).
Proceedings of the California Academy of Sciences 35:1 29-1 39.
Shook, G. A. 1 984. Checklist of tiger beetles from Idaho (Coleoptera: Cicindelidae). Great
Basin Naturalist 44:1 59-1 60.
Scheaffer, R. L., W. Mendenhall, and L Ott. 1 979. Elementary survey sampling. Duxbury
Press, North Scitvate, Massachusetts.
ER'S CARD
2 no. 94-1
s tiger beetle
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QL 84.2 .L352 no. 94-1
88055211
Bruneau Dunes tiger beetle
inventory
DUV1 LlbHAHY
BLDG50, ST-150A
DENVER FEDERAL CENTER
P.O. BOX 25047
DENVER, COLORADO 80225
BUREAU OF LAND MANAGEMENT
Idaho State Office
3380 Americana Terrace
Boise, Idaho 83706
BLM-ID-PT-94-0084050