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**Butterfly Monitoring Protocol for Four Prairie Parks**

**Northern Prairie Wildlife Research Center Inventory and Monitoring
Protocol**

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U.S. Department of the Interior**

**U.S. Geological Survey**

**REVIEW NOTICE**

This report has been prepared for peer-review through the procedures of
the U.S. Geological Survey, Northern Prairie Wildlife Research Center.
Mention of trade names or commercial products does not constitute
endorsement or recommendation for use.

U.S. Department of the Interior

U.S. Geological Survey

**Butterfly Monitoring Protocol for Four Prairie Parks**

by

Diane Debinski^1^

Scott Mahady^1^

William M. Rizzo^2^

Gary D. Willson^2^

^1^Department of Animal Ecology

124 Science II

Iowa State University

Ames, Iowa 50011

^2^U.S. Geological Survey

Northern Prairie Wildlife Research Center

Missouri Project Office

302 Gentry Hall

University of Missouri-Columbia

Columbia, Missouri 65211

Prepared for Great Plains Prairie Cluster Long-Term Ecological
Monitoring Program

National Park Service

August 2000

**ACKNOWLEDGMENTS**

Funding for this project was provided through the Inventory and
Monitoring Program of the U.S. Geological Survey and the National Park
Service. We thank Robert Cox, Ralph Grundel, Ron Royer, and an anonymous
reviewer for the National Park Service for their constructive comments
on an earlier draft of this protocol. Thanks also are due to Deanna
Adkins who edited the final version of the protocol.

**PREFACE**

The original draft of the Butterfly Monitoring Protocol for Four Prairie
Parks was synthesized by Drs. William M. Rizzo and Gary D. Willson from
the completion reports on the butterfly species inventories in the four
parks prepared by Mr. Scott Mahady and Dr. Diane Debinski and from the
completed thesis of Mr. Mahady.

**CONTENTS**

**Page**

**ACKNOWLEDGMENTS** iv

**PREFACE** v

**LIST OF TABLES** viii

**LIST OF FIGURES** ix

**1.0 INTRODUCTION**

> 1.1 Background 1
>
> 1.2 Objectives of butterfly monitoring 1

**2.0** **PROTOCOL DESIGN**

> 2.1 Sampling methods 2
>
> 2.2 Sampling sites 2
>
> 2.3 Sampling intensity 3
>
> 2.4 Sampling frequency and timing 3

**3.0 FIELD IMPLEMENTATION**

> 3.1 Field sampling 4

3.2 Additional field data collection 5

**4.0** **CALCULATION OF METRICS AND STATISTICAL TESTING**

> 4.1 Butterfly metrics by guild and habitat type 5
>
> 4.2 Long-term data analysis 6

**5.0 DATA MANAGEMENT**

> 5.1 Database design 6
>
> 5.2 Data entry and checking 7
>
> 5.3 Annual reports 8

**6.0 REFERENCES** 9

**APPENDIX** **A**

Locations of butterfly monitoring transects in three parks of the
Prairie

> Cluster Program 24

**APPENDIX** **B**

Annual report on butterfly monitoring at Homestead National Monument of
America 25

**LIST OF TABLES**

**Table Page**

1\. Habitat types and number of sampling transects per habitat in four
parks of the

> Prairie Cluster Program 13

2.  Recommended temporal sampling windows in four parks of the

> Prairie Cluster Program 14

3.  Equipment and supplies necessary for butterfly monitoring 15

4.  Field data sheet for recording butterfly count data and weather data
    16

5.  Butterfly species by guild in four prairie parks and adjacent
    prairies

> during 1997 and 1998 18

6\. An example of an Access database and its tables and linkages for
storage and

> handling of the butterfly monitoring data 22

**LIST OF FIGURES**

**Figure Page**

1\. Number of butterfly species recorded vs. number of transects sampled

> for six prairies 23

**1.0 INTRODUCTION**

1.1 [Background]{.underline}

Habitat loss and fragmentation are two of the primary factors leading to
species extinctions during this century. Tallgrass prairie habitat in
the Midwestern United States has been particularly affected. Only about
1% of the original tallgrass prairie ecosystem still remains (Swengel
1996). The drastic loss of prairie habitat, the disruption of natural
disturbance regimes and nutrient cycles, and the isolation of the
remaining tallgrass prairie habitat makes the preservation of
prairie-dependent species a unique challenge (Johnson and Simberloff
1974; Leach and Givnish 1996; Collins et al. 1998; Kaiser 1998; Schlicht
and Orwig 1998).

The butterfly communities of tallgrass prairie remnants can be viewed as
indicators of ecosystem health, much like aquatic macroinvertebrates
have long been used as indicators of water quality. In addition, the
butterfly communities can serve as potential indicators for impacts on a
broader spectrum of invertebrate species (New et al. 1995). As an
indicator group, butterfly communities offer several advantages: 1)
Their habitat preferences and host plant associations are relatively
well-known, which allows classification of individual species into
guilds (Sharp et al. 1974; Scott 1986; Opler and Malikul 1992; Panzer et
al. 1995; Swengel 1996; 1998); 2) Much literature exists on
metapopulation dynamics, dispersal, and the effects of different
habitats on butterfly movement behavior (Sharp et al. 1974; Fahrig and
Paloheimo 1987; Hanski et al. 1995; Hanski et al. 1996; Hill et al.
1996; Neve et al. 1996; Peterson 1997; Haddad 1999); and 3) Obligate
prairie species respond quickly to changes in native vegetation (Miller
and Harris 1977; Saunders et al. 1991).

While this protocol was developed to monitor the butterfly community to
determine the status of the ecosystem health of the existing habitat
types, butterfly monitoring also could be used to evaluate the impact of
management practices or extraordinary natural events. These could
include evaluations of practices such as prescribed fire, mowing,
herbicide application, or grazing, or impacts caused by natural events
such as flooding or invasion of exotic species.

With the growing emphasis on ecosystem management, the National Park
Service (NPS) must possess protocols for monitoring the status of
butterfly communities within an ecosystems context. This protocol was
developed for use by four of the national parks within the Great Plains
Prairie Cluster Long-Term Ecological Monitoring Program (hereafter
referred to as the Prairie Cluster Program). Two of the four parks,
Pipestone National Monument (PIPE) in Minnesota, and Homestead National
Monument of America (HOME) in Nebraska have both restored and native
tallgrass prairie. The other two parks, Wilson's Creek National
Battlefield (WICR) in Missouri, and Effigy Mounds National Monument
(EFMO) in Iowa, have restored and native tallgrass prairie and oak
savanna/woodland. WICR also has glade habitats.

1.2 [Objectives of butterfly monitoring]{.underline}

Butterfly communities were chosen to serve as indicators of ecosystem
health for prairie, savanna/woodland and glade habitats. Butterfly
monitoring allows park managers to detect and describe long-term changes
in the butterfly communities associated with these habitats.
Specifically, the monitoring objectives are: 1) to quantify temporal
changes in the species composition, abundance, and richness of butterfly
guilds for selected habitats in each park; and 2) to quantify
differences in the species composition, abundance, and richness of
butterfly guilds among park habitats. The spatial analyses also can be
used by park managers to assess the effectiveness of management
practices such as prescribed fire or to evaluate the success of restored
prairie compared to native prairie remnants.

**2.0 PROTOCOL** **DESIGN**

2.1 [Sampling methods]{.underline}

This protocol basically follows the procedures described in Mahady
(1999), which are modified from the transect technique described by
Pollard (1977). Unintrusive transect surveys such as the "Pollard walk"
are adequate and efficient for monitoring butterfly communities,
especially in protected areas such as national parks where rare species
are known to occur (Murphy 1988). Furthermore, the fixed transects of
Pollard walk counts are uniform with respect to area covered and time
spent, which allows more rigorous statistical analysis (Royer et al.
1998). The fixed nature of the transects also allows for concurrent
monitoring of other natural resources such as the plant community.
Pollard transects typically use count data to monitor abundance of
butterfly species (Swengel 1996) and species richness (Mahady 1999).

2.2 [Sampling sites]{.underline}

Each park unit was sampled in 1997 and 1998 to inventory the butterfly
community and propose management recommendations to the resource
managers. These parks were chosen for butterfly monitoring because
prairie plant community studies within fragmented habitats also were
ongoing in these parks. Within each park, sampling was carried out along
transects in different habitat types, which varied among the different
parks. The habitat types sampled in each park are shown in Table 1
(Mahady and Debinski 1999a; 1999b; 1999c; 1999d). Each transect is 5-m
wide by 50-m long. The transect length was established for conformity
with vegetation sampling (Buck et al. 2000; Anonymous 1993) so that
possible future analyses of butterfly/plant relationships could be made.

In 1998, where possible, 6 transects were placed in each habitat
parallel to previously established plant community transects which had
been located randomly (Mahady 1999). In 1999, two additional transects
were sampled in some prairies large enough to accomodate them (Mahady
1999). Subsequent analyses (see section 2.3) indicated that six
transects are sufficient for sampling prairies the size of those listed
in Table 1. All available transect locations are shown in Appendix A.
Transect locations for EFMO are not available.

In prairies larger than 4 ha, butterfly transects were located at least
50 m apart. In smaller prairies, the size and shape of the habitat
dictated the placement of transects. In these areas, the transects were
located to maximize both the distance between transects and the distance
from the habitat edge. Transects occasionally had to be curvilinear to
be accommodated within a habitat type. In an extreme case (North Bloody
Hill at WICR), the habitat was so fragmented that the transects
consisted of sampling the entire area. In Appendix A, transect locations
for this site are actually count points. In a number of areas, 6 or
fewer 50 m transects were all that could be accommodated within the
habitat (native prairie at HOME; all sites at EFMO). The number of
transects recommended for sampling in each park and habitat is shown in
Table 1.

2.3 [Sampling intensity]{.underline}

Some of the data collected by Mahady (1999) were analyzed to assess the
adequacy of the number of transects. The number of species from one
transect randomly selected from the dataset was used to represent
"transect 1\" for a given sampling date. A second randomly selected
transect was added to the first transect to represent "transects 1 and
2" (i.e. sampling without replacement). This process was repeated until
all the data from the 6 transects were recorded. This process was
undertaken 20 times, and the mean results are shown in Figure 1 for
several sites. These results showed that for parks similar in size to
most of those used in the development of this protocol, 85% of the taxa
present were recorded by sampling only 3 to 4 transects, so 6, 50-m
transects were more than adequate for this sampling effort. These
results also suggest that for very large parks such as Tallgrass Prairie
National Preserve, 6 transects are not adequate. For larger parks, 8 or
more transects should be initially established, and their adequacy
should be evaluated by a method similar to that described above.

2.4 [Sampling frequency and timing]{.underline}

Emergence periods (i.e. flights) for some butterfly species are brief,
but individuals may be very abundant. For instance, Mahady (personal
communication) reported 101 sightings of the powesheik skipper at PIPE
on 27 June 1998, but none on 26 June 1997, probably because the 1997
flight was slightly earlier. To increase the likelihood of capturing
such ephemeral flights, especially for species with different emergence
dates during the year, transects should be sampled four times during the
growing season. This doubles the sampling effort used by Mahady and
Debinski (1999a; 1999b; 1999c; 1999d), but the objectives of those
studies were different than the goals of this monitoring protocol. The
sampling windows suggested for the various parks are shown in Table 2.
These windows were selected based on previous sampling experience
(Mahady and Debinski 1999a; 1999b; 1999c; 1999d) and analysis of the
typical emergence times of butterfly species (Richard and Heitzman
1987).

Certain minimum weather conditions also are required for an adequate
sample. Butterfly flight activity is decreased by cool temperatures or
heavy cloud cover. Cloud cover is not as important a limitation as
temperature, so the percentage of cloud cover can become relatively high
and not limit butterfly activity if the temperature also is high. The
temperature must exceed 18°C, and cloud cover must be less than 70%. If
cloud cover exceeds 70%, the temperature must exceed 23°C. Surveys must
be conducted on days when wind gusts are less than 45 kph (30 mph) or on
days with prevailing winds less than 30 kph (20 mph). There are no
standard guidelines for sampling constraints, but the limits recommended
here are similar to those used by other butterfly researchers (Pollard
1977; Ron Royer, personal communication, uses 50% cloud cover, 16°C, and
Beaufort scale \< 4); (i.e. \< 20 km h^-1^), and their continued use
will ensure consistency in the physical conditions for data collection.
Surveys are conducted between 0900 and 1730 hrs, because butterfly
activity is low before 0900 due to cooler temperatures.

Temperatures are determined using a stem field thermometer placed in the
shade under vegetation. This will underestimate actual temperature, and,
therefore, is a conservative approach to sampling during periods of
butterfly activity. Cloud cover is a visual estimate of the percentage
of clouds obscuring the sky, excluding the lower 30° from the horizon
upward on all sides of the observer (resulting in a total of 120° of sky
for the estimate). Wind speeds are determined in the field with a wind
meter (e.g. Wind Wizard®).

**3.0 FIELD IMPLEMENTATION**

3.1 [Field sampling]{.underline}

A list of supplies and equipment used for butterfly monitoring is given
in Table 3. Prior to sampling (the day prior to sampling at the latest),
the transects should be flagged every 10 m along each boundary of the
5-m transect width. The observer then walks down the middle of the
transect. The 2.5-m distance on either side of the observer is the
maximum distance for reliable visual identification (Pollard 1977). All
butterflies are identified by a single observer walking at a constant
pace of 10-m per minute. A stopwatch is used to ensure the correct speed
is followed. The timing is stopped whenever additional time is needed
for capture and identification of some individuals or when large numbers
of individuals are encountered. The observer identifies all individuals
to species except for individuals of the genus *Phyciodes*, which are
difficult to identify to species in the field. Extra care also should be
taken in identifying species of the genera *Erynnis* and *Thorybes*, but
they can be adequately identified to species in the field. Individual
butterflies that cannot be identified in flight are captured using a
standard butterfly net and then identified. The number of individuals of
each species encountered during each count is recorded on a data sheet
(Table 4) along with the number of counts made on each transect. The
example in Table 4 includes the entire park species list by guild
(Mahady and Debinski 1999c). The observer also should carry the entire
species list compiled by Mahady and Debinski (1999a; 1999b; 1999c;
1999d) for aid in identifying new species. In addition to the entire
species list for the four parks, this list also contains species that
were found in prairies outside the parks and that would likely be
encountered in future sampling within the parks.

Each transect is surveyed 6 times during a sampling visit for a total
walking time of 30 minutes at the required pace. A minimum of 15 minutes
should elapse between counts to allow the butterfly community to recover
from the disturbance of the previous sample. If there are other habitat
transects nearby, these can be sampled during the waiting period. Since
the total walking time for a transect is just 30 minutes, the waiting
period increases the temporal span of the counts to 2 hrs, which allows
for incorporation of some diel effects on the activity of different
butterfly species. Repeated surveys, rather than a single extended
count, are recommended because uncommon species were occasionally
tallied during later counts (Mahady, personal observations) and because
the actual identification and enumeration time is extended as needed
when the observer is not walking.

Butterfly species respond differently to the effects of sampling
disturbance. Some species will be largely unaffected, yet others may be
driven away. However, because these behaviors can be assumed to be
constant from year to year and because the focus of the protocol is to
detect changes over time, any sampling bias introduced by disturbance
should not affect the outcome of the protocol sampling objectives.
Sampling should be undertaken by personnel with advanced training (e.g.
Master's degree) in entomology with special training in butterfly
taxonomy. The field guides with the greatest relevance to these parks
are Richard and Heitzman (1987) and Glassberg (1999).

3.2 [Additional field data collection]{.underline}

The physical field conditions under which the butterfly counts are made
also are recorded on the data sheet (Table 4) in order to assess the
possibility that weather conditions may have affected survey results.
Air temperature (^°^C), wind speed (kph), wind gusts (kph), and cloud
cover (%) should be recorded prior to sampling and again after sampling
is completed.

Butterfly population dynamics are intimately tied to plant community
dynamics. While there may be important interannual differences in plant
phenology (e.g. flowering time), plant community data collected to date
indicate that the plant community is composed primarily of perennial
species that show few interannual differences in community composition
(DeBacker et al. 1998; DeBacker 1999). However, long-term successional
changes are likely in disturbed habitats. Park managers interested in
maintaining native plant diversity must also understand the status and
trends of prairie plant communities. The Prairie Cluster Program is
nearing completion of a protocol for monitoring plant communities. This
protocol describes the collection of percent foliar cover of all
herbaceous and shrub species and basal area for trees using fixed plots
sampled in late spring/early summer and late summer. Metrics such as
Shannon diversity and species richness then are calculated from the data
to allow detection of changes from year to year and from a baseline year
(Buck et al. 2000). Plant community monitoring programs have been
established at the parks used for butterfly sampling. Other parks or
areas wishing to monitor butterfly communities also should establish a
plant community monitoring program. A weather monitoring program (e.g.,
Akyuz et al. 2000) also can provide valuable insights into butterfly
community dynamics.

4.0 **CALCULATION OF METRICS AND STATISTICAL TESTING**

4.1 [Butterfly metrics by guild and habitat type]{.underline}

Each butterfly species in the baseline dataset is tabulated by
ecological guild in Table 5. The classification of butterfly species as
prairie obligates is based on a large body of research (Scott 1986;
Opler and Malikul 1992; Panzer et al. 1995; Swengel 1996; Schlict and
Orwig 1998). The relative dependence of other butterfly species on
particular habitats is less well-known and was derived from general
descriptions of the species (Scott 1986; Opler and Malikul 1992).

The total abundance of the species comprising each guild can be tested
among years or time periods using repeated-measures analysis of variance
and multiple range testing (e.g., Duncan's test) because transects are
permanently fixed. Comparison of different sites also may be carried out
using analysis of variance. The data should first be tested for
compliance to the assumptions of normality and homoscedasticity required
for parametric analysis of variance (SAS Institute Inc. 1989-1996). Data
transformations (e.g., log, rank, square root) may be required. If the
data cannot be adequately transformed, non-parametric analyses analogs
of analysis of variance, such as the Kruskal-Wallis test, can be used.
Non-parametric tests are less sensitive to data outliers, which greatly
contribute to heteroscedasticity. High variability can occur in
butterfly surveys due to low abundance values, infrequent occurrence
(i.e., locally rare species), or patchy distributions.

Species richness is simply the sum of the number of species, by guild,
recorded for a transect on a given date and habitat. High butterfly
species richness is indicative of high quality habitat, i.e. habitats
with high vegetative diversity. Species richness also can be used to
assess changes at a site over time or to compare the richness of
restored prairies to native prairies (Selser 1992). Species richness
also is analyzed by the procedures used for analyzing the abundance
data. In addition, an easy way to compare the species richness in pairs
of habitats, or pairs of time periods, is through the use of similarity
coefficients such as the Jaccard index (Smith 1990).

It is calculated as:

SC~J~ = c/(A + B - c),

where c equals the number of species found in both habitats, A equals
the total number of species in habitat A, and B is the total number of
species in habitat B. Values range from 0 (no species common to both) to
1 (all species occur in both habitats). Thus, while richness may
indicate good habitat diversity, it could be possible that the butterfly
community composition in two equally diverse areas arises from totally
different community composition. Use of similarity indices can thus
augment the species richness information by offering a way to evaluate a
data set in reference to a desired target (i.e., a native prairie
butterfly community).

4.2 [Long-term data analyses]{.underline}

Ultimately, the goal of monitoring efforts should be the detection of
changes in community attributes over time. For example, one might wish
to detect a 20% change in abundance with 90% confidence. It is intuitive
that changes should be detectable before changes become great enough to
result in adverse impacts. However, the magnitude of these changes
causing adverse impacts are largely unknown. In addition, efforts to
assess sampling adequacy, (e.g., power analysis) need to be based on
enough years of data to incorporate the full range of interannual
variability of the parameter. Because butterfly abundances may change
several orders of magnitude between years, it is highly unlikely that
application of techniques such as power analysis to two years of
available data (Mahady 1999) is likely to result in a useful
recommendation of sampling effort. A more reasonable approach would be
to continue butterfly monitoring and address this question when
additional years of data are available for analysis. A minimum of 5
years of annual data should be collected and analyzed to assess sampling
adequacy. However, a final consideration is that techniques for
estimating required sampling effort may not be very realistic in an
ecological sense. In a number of cases within these four parks,
essentially the entire habitat was surveyed so that additional transects
could not be added regardless of the results of power analysis.

**5.0 DATA MANAGEMENT**

5.1 [Database design]{.underline}

Original data sheets with the butterfly count data and the data on the
physical variables should be stored at the Prairie Cluster Program
office. If a contractor carries out the monitoring, the contractor
should keep a copy of the original data sheets.

The Prairie Cluster Program office uses a Microsoft® Access database to
store the long-term archive, though other statistical or spreadsheet
programs also can be used. If other storage media are used, however, it
is important for continuity or "institutional memory" that metadata
descriptions be incorporated into the data-storage process. The data
within the Access database cannot serve as a direct medium for data
input into statistical packages. The database must be queried to compile
a table of the required data, which is exported into a suitable
spreadsheet format that then serves as the input for the statistical
package.

The Access butterfly database is designed for compatibility with other
Prairie Cluster Program monitoring databases, so it incorporates
standardized site and event information. The database comprises 6 tables
in a relational design (Table 6). Tbl_Butterfly_Data contains the
abundance (count) data by species, with alphanumeric codes identifying
site, sampling event (date), and species information. Related site,
event, taxa, and weather (field data) tables are associated with
Tbl_Butterfly_Data through "one-to-many" controlled links. These
linkages impose referential integrity between the core data and related
information, ensure data quality, and provide for easy incorporation of
data updates. For example, only after all sample event and investigator
data have been entered could a particular sample event for data entry be
selected. Additionally, the database linkages can speed data entry. For
instance, a species guild assignment is updated in a single location in
Tbl_Butterfly_Species rather than for each species (potentially
hundreds).

The data stored in Tab_Site, Tbl_Event, Tbl_Butterfly_Species,
Tbl_Butterfly_Guilds, and Tbl_Weather are shown in Table 6. Tbl_Site
includes UTM coordinates for the sampling sites, where available, and
other descriptive information. Tbl_Event includes sampling dates,
investigator names, and field notes. Tbl_Weather contains the physical
field data collected along with the butterfly abundance data.
Tbl_Butterfly_Species includes scientific names, guild assignments, and
taxon codes. Tbl_Butterfly_Guilds contains butterfly guilds
descriptions. A 6-letter code is assigned to each butterfly taxon during
data entry. It consists of the first 3 letters of the genus name and the
first 3 letters of the species name. In case of a duplicate code for a
new species, the first letter of the species name resulting in a
different code is used. Taxon codes are necessary for data manipulation
in many spreadsheet and statistical packages that do not allow long
variable names.

5.2 [Data entry and checking]{.underline}

The database must be updated as new data are acquired, which can result
in significant numbers of transcription errors. To reduce transcription
errors, data entry is accomplished by using customized forms that
prevent entry of duplicate or incorrect taxonomic records. There is a
unique identification value for each site/date combination. Most other
variables can be selected from drop-down menus. Each species entry is
selected from drop-down menus and linked to the Tbl_Taxa so that only
valid species names can be entered into the database, which eliminates
typographic mistakes. If a species name is not accepted, it is probably
either a species new to the database or the result of a nomenclature
change. A nomenclature update form is attached via a button to allow
quick checking for the current accepted name. If a species is not
presently included in the taxa table, a form is available to update this
table with the new attribute data.

Data verification occurs immediately after data entry by checking the
accuracy of the computerized records against the original sources.
Someone familiar with butterfly taxonomy should do the manual
verification. In addition, database queries can help validate the
accuracy and completeness of the database after all the season's data
entry has been made. The query should search records with a site code
from one park and an event code from a different park.

5.3 [Annual reports]{.underline}

An annual report should be prepared giving the results from each park
for each year of sampling. The report should present the results of the
abundance and species richness analyses for the butterfly guilds for
each park habitat. Reports also should describe the physical conditions
under which the data were collected. An example report is given in
Appendix B.

**6.0 REFERENCES**

Akyuz, F. A., P. Guinan, and G. D. Willson. 2000. Weather monitoring
protocol for two prairie parks. U.S. Geological Survey, Biological
Resources Division, Northern Prairie Wildlife Research Center, Missouri
Field Station, 302 Gentry Hall, University of Missouri-Columbia,
Columbia, MO 65211.

\[Anonymous\]. 1993. Methods manual for Konza Prairie Research Natural
Area. Manhattan: Kansas State University.

Buck, C., G. D. Willson, L. Thomas, M. DeBacker, and W. M. Rizzo. 2000.
Draft plant community monitoring protocol for six prairie parks. U.S.
Geological Survey, Biological Resources Division, Northern Prairie
Wildlife Research Center, Missouri Field Station, 302 Gentry Hall,
University of Missouri-Columbia, Columbia, MO 65211.

Collins, S. L., A. K. Knapp, J. M. Briggs, J. M. Blair, and E. M.
Steinauer. 1998. Modulation of diversity by grazing and mowing in native
tallgrass prairie. Science 280:745-747.

DeBacker, M. 1999. 1998 plant community monitoring for Wilson's Creek
National Battlefield. Final Report to the Prairie Cluster Long-Term
Ecological Monitoring Program, National Park Service.

DeBacker, M., L. Thomas, and K. Grabner. 1998. 1997 plant community
monitoring for Effigy mounds National Monument, Pipestone National
Monument, Scotts Bluff National Monument, Wilson's Creek National
Battlefield. Final Report to the Prairie Cluster Long-Term Ecological
Monitoring Program, National Park Service.

Fahrig, L. and J. E. Paloheimo. 1987. Interpatch dispersal of the
cabbage butterfly. Canadian Journal of Zoology 65:616-622.

Glassberg, J. 1999. Butterflies through Binoculars. The East. New York:
Oxford University Press.

Haddad, N. M. 1999. Corridor use predicted from behaviors at habitat
boundaries. American Naturalist 153:215-227.

Hanski, I., A. Moilanen, T. Pakkala, and M. Kuussaari. 1996. The
quantitative incidence function model and persistence of an endangered
butterfly metapopulation. Conservation Biology 10:578-590.

Hanski, I., T. Pakkala, M. Kuussaari, and G. Lei. 1995. Metapopulation
persistence of an endangered butterfly in a fragmented landscape. Oikos
72:21-28.

Hill, J. K., C. D. Thomas, and O. T. Lewis. 1996. Effects of habitat
patch size and isolation on dispersal by hesperia-comma butterflies -
implications for metapopulation structure. Journal of Animal Ecology
65:725-735.

Johnson, M. P. and D. S. Simberloff. 1974. Environmental determinants of
island species numbers in the British Isles. Journal of Biogeography
1:149-154.

Kaiser, J. 1998. Ecology - bison prime prairie biodiversity. Science
280:677.

Leach, M. K. and T. J. Givnish. 1996. Ecological determinants of species
loss in remnant prairies. Science 273:1555-1558.

Mahady, S. 1999. Conservation of tallgrass prairie butterfly species in
a highly fragmented landscape. M.Sc. Thesis. Iowa State University.
Ames, IA.

Mahady, S. and D. Debinski. 1999a. Habitat management plan for butterfly
species at Effigy Mounds National Monument. A report to the Missouri
Field Station, U. S. Geological Survey. Iowa State University, Ames, IA.

Mahady, S. and D. Debinski. 1999b. Habitat management plan for butterfly
species at Homestead National Monument. A report to the Missouri Field
Station,

U. S. Geological Survey. Iowa State University, Ames, IA.

Mahady, S. and D. Debinski. 1999c. Habitat management plan for butterfly
species at Pipestone National Monument. A report to the Missouri Field
Station, U. S. Geological Survey. Iowa State University, Ames, IA.

Mahady, S. and D. Debinski. 1999d. Habitat management plan for butterfly
species at Wilson's Creek National Battlefield. A report to the Missouri
Field Station, U. S. Geological Survey. Iowa State University, Ames, IA.

Miller, R. I. and L. D. Harris. 1977. Isolation and extirpations in
wildlife reserves. Biological Conservation 12:311-315.

Murphy, D. D. 1988. Opinion: are we studying our endangered butterflies
to death? Journal of Research on the Lepidoptera 26:236-239

Neve, G., L. Mousson, and M. Baguette. 1996. Adult dispersal and
genetic-structure of butterfly populations in a fragmented landscape.
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New, T. R., R. M. Pyle, J. A. Thomas, C. D. Thomas, and P. C. Hammond.
1995. Butterfly conservation management. Annual Review of Entomology
40:57-83.

Opler, P. A. and V. Malikul. 1992. A Field Guide to Eastern Butterflies.
Boston: Houghton Mifflin Co.

Panzer, R., D. Stillwaugh, R. Gnaedinger, and G. Derkovitz. 1995.
Prevalence of remnant dependence among prairie and savanna inhabiting
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Peterson, M. A. 1997. Host-plant phenology and butterfly dispersal -
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Pollard, E. 1977. A method for assessing changes in the abundance of
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Missouri Department of Conservation. Jefferson City, MO.

Royer, R. A., J. E. Austin, and W. E. Newton. 1998. Checklist and
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Cary, NC.

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University Press.

Selser, E. J. 1992. Diversity, abundance, and distribution of butterfly
species in remnant and tallgrass prairie. In: Wickett, R. G., Lewis, P.
D., Woodliffe, A., and Pratt, P. editors. Proceedings of the Thirteenth
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prairie butterflies. Biological Conservation 76:73-85.

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tallgrass prairie and pine-barrens. Biological Conservation 83:77-89.

Table 1. Habitat types and recommended number of sampling transects per
habitat in four parks of the Prairie Cluster Program. \* indicates sites
containing the maximum possible number of [transects.]{.underline}

Park/Habitat Number of transects

Effigy Mounds National Monument

> Restored prairie 2\*
>
> Goat prairie 6\*

Homestead National Monument of America

> Restored prairie 6\*
>
> Schoolhouse native prairie 6\*

Pipestone National Monument

> Restored prairie 6
>
> Native prairie 6
>
> Native prairie with rare orchids 2\*

Wilson's Creek National Battlefield

> Bloody Hill (glade/woodland) 6
>
> North Bloody Hill (glade/woodland/grassland) surveyed in its entirety
>
> Upland savanna 4\*
>
> Ray House restored prairie 2\*

[Table 2. Recommended temporal sampling windows in four parks of the
Prairie Cluster Program.]{.underline}

Effigy Mounds National Monument

> May 12-17 June 15-25 July 2-8 August 1-6

Homestead National Monument of America

> May 3-9 June 7-15 July 20-28 August 18-24

Pipestone National Monument

> May 18-26 June 26-July 2 July 8-13 August 7-16

Wilson's Creek National Battlefield

> April 4-30 May 28-June 5 July 13-18 August 26-31

[Table 3. Equipment and supplies necessary for butterfly
monitoring.]{.underline}

For laying out transects:

tape measure

flags

For butterfly counting and identification:

46 cm (18 in) diameter aerial net (e.g. from Bioquip®)

park species list

total species list for all parks

data sheets

pencils

field guides

stopwatch

For additional data collection:

wind meter (e.g. Wind Wizard ®)

stem thermometer

[Table 4. Field data sheet for recording butterfly count data and
weather data.]{.underline}

Date: [1 June 1999]{.underline} Park name: [Pipestone NM]{.underline}

Start time: [1000]{.underline}

Temperature: [20 ^°^C]{.underline} Wind speed: [5 kph]{.underline} Wind
gusts: [10 kph]{.underline} Cloud cover: [10%]{.underline}

End Time: [1400]{.underline}

Temperature: [23 ^°^C]{.underline} Wind speed: [10 kph]{.underline} Wind
gusts: [15 kph]{.underline} Cloud cover: [30%]{.underline}

Site/habitat type: [Restored prairie]{.underline} Surveryor(s):

[Transect: 1 2 3 4 5 6]{.underline}

[Counts:]{.underline}

[Species Common name Taxon code Number of individuals]{.underline}

**Open habitat generalist species**

[*Colias eurytheme* alfalfa butterfly COLEUR]{.underline}

[*Colias philodice* clouded sulfur COLPHI]{.underline}

[*Danaus plexippus* monarch DANPLE]{.underline}

[*Euptoieta claudia* variegated fritillary EUPCLA]{.underline}

[*Everes comyntas* eastern-tailed blue EVECOM]{.underline}

[*Hemiargus isola* Mexican blue HEMISO]{.underline}

[*Phyciodes tharos* pearl crescent PHYTHA]{.underline}

[*Pieris rapae* cabbage PIERAP]{.underline}

[*Vanessa atalanta* red admiral VANATA]{.underline}

[*Vanessa cardui* painted lady VANCAR]{.underline}

[*Wallengrenia egeremet* broken dash WALEGE]{.underline}

Table 4. Continued**.**

[Transect: 1 2 3 4 5 6]{.underline}

[Counts:]{.underline}

[Species Common name Taxon code Number of individuals]{.underline}

**Grassland specialist species**

[*Coenonympha tullia* ringlet COETUL]{.underline}

[*Papilio polyxenes* black swallowtail PAPPOL]{.underline}

[*Plebejus melissa* melissa blue PLEMEL]{.underline}

[*Polites themistocles* tawny edged skipper POLTHE]{.underline}

[*Speyeria atlantis* atlantis fritillary SPEATL]{.underline}

**Prairie obligate species**

[*Oarisma powesheik* powesheik skipper OARPOW]{.underline}

[*Speyeria aphrodite* aphrodite fritillary SPEAPH]{.underline}

[*Speyeria idalia* regal fritillary SPEIDA]{.underline}

**Woodland species**

[*Ancyloxpha numitor* least skipperling ANCNUM]{.underline}

[*Cercyonis pegala* common wood nymph CERPEG]{.underline}

[*Euphyes vestris* dun skipper EUPVES]{.underline}

[*Papilio glaucus* tiger swallowtail PAPGLA]{.underline}

**Wetland specialist species**

[*Lethe eurydice* northern pearl eye LETEUR]{.underline}

[*Polites mystic* long dash POLMYS]{.underline}

Table 5. Butterfly species by guild in four prairie parks and adjacent
prairies during 1997 and 1998. WICR = Wilson's Creek National
Battlefield, HOME = Homestead National Monument of America, EFMO =
Effigy Mounds National Monument, and PIPE = Pipestone National Monument.
Species with no park indicated were recorded during sampling of other
nearby prairies (Mahady 1999). Guilds are coded as HG = open habitat
generalist species, GS = grassland specialist species, PO = prairie
obligate species, WS = woodland [species, WE = wetland specialist
species, and SS = savanna specialist species.]{.underline}

[Species Common name Taxon code Parks where present]{.underline} **Open
habitat generalist species**

*Atalopedes campestris* sachem ATACAM WICR

*Celastrina argiolus* spring azure CELARG WICR HOME EFMO

*Chlosyne nycteis* silvery checkerspot CHLNYC WICR

*Colias eurytheme* alfalfa butterfly COLEUR PIPE WICR HOME

*Colias philodice* clouded sulfur COLPHI PIPE WICR HOME EFMO

*Danaus plexippus* monarch DANPLE PIPE WICR HOME EFMO

*Euptoieta claudia* variegated fritillary EUPCLA PIPE WICR HOME

*Everes comyntas* eastern-tailed blue EVECOM PIPE WICR HOME

*Hemiargus isola* Mexican blue HEMISO PIPE WICR HOME

*Hylephila phyleus* fiery skipper HYLPHY

*Leptotes cassius* tropical striped blue LEPCAS

*Leptotes marina* striped blue LEPMAR

*Limenitis archippus* viceroy LIMARC EFMO

*Nymphalis antiopa* mourning cloak NYMANT EFMO

*Pholisora cattulus* common sooty wing PHOCAT WICR

*Phyciodes tharos* pearl crescent PHYTHA PIPE WICR HOME EFMO

*Pieris rapae* cabbage PIERAP PIPE WICR HOME

*Precis coenia* buckeye PRECOE WICR EFMO

*Pyrgus communis* checkered skipper PYRCOM HOME

*Speyeria cybele* great spangled fritillary SPECYB WICR HOME EFMO

*Strymon melinus* grey hairstreak STRMEL WICR HOME EFMO

*Vanessa atalanta* red admiral VANATA PIPE WICR HOME EFMO

*Vanessa cardui* painted lady VANCAR PIPE HOME

Table 5. Continued.

[Species Common name Taxon code Parks where present]{.underline} **Open
habitat generalist species (continued)**

*Vanessa virginiensis* American painted lady VANVIR WICR EFMO

*Wallengrenia egeremet* broken dash WALEGE PIPE WICR EFMO

**Grassland specialist species**

*Atrytone logan* Delaware skipper ATRLOG WICR HOME EFMO

*Chlosyne gorgone* gorgone checkerspot CHLGOR HOME

*Coenonympha tullia* ringlet COETUL PIPE

*Colias cesonia* dog face COLCES

*Erynnis martialis* mottled dusky wing ERYMAR HOME

*Eurema lisa* little sulfur EURLIS WICR HOME EFMO

*Eurema niccipe* sleepy orange EURNIC WICR

*Harkenclenus titus* coral hairstreak HARTIT WICR

*Lycaena hyllus* bronze copper LYCHYL HOME

*Nastra lherminier* swarthy skipper NASLHE WICR

*Papilio polyxenes* black swallowtail PAPPOL PIPE WICR

*Phoebis sennae* cloudless sulfur PHOSEN WICR HOME

*Pieris protodice* checkered white PIEPRO HOME

*Plebejus melissa* Melissa blue PLEMEL PIPE

*Polites peckius* Peck's skipper POLPEC WICR HOME

*Polites themistocles* tawny edged skipper POLTHE PIPE WICR HOME

*Speyeria atlantis* atlantis fritillary SPEATL PIPE

*Thorybes bathyllus* southern cloudy wing THOBAT WICR

**Prairie obligate species**

*Atrytonopsis hianna* dusted skipper ATRHIA

*Hesperia ottoe* prairie skipper HESOTT

Table 5. Continued.

[Species Common name Taxon code Parks where present]{.underline}
**Prairie obligate species (continued)**

*Polites origenes* crossline skipper POLORI

*Boloria bellona* meadow fritillary BOLBEL

*Hesperia dacotae* Dakota skipper HESDAC

*Oarisma powesheik* powesheik skipper OARPOW PIPE

*Problema byssus* byssus skipper PROBYS EFMO

*Speyeria aphrodite* aphrodite fritillary SPEAPH PIPE EFMO

*Speyeria idalia* regal fritillary SPEIDA PIPE WICR HOME EFMO

*Hesperia leonardus* Leonard's skipper HESLEO EFMO

**Woodland species**

*Achalarus lyciades* hoary edge ACHLYC

*Anea andria* goatweed ANEAND WICR

*Ancyloxpha numitor* least skipperling ANCNUM PIPE

*Asterocampa celtis* hackberry ASTCEL WICR EFMO

*Asterocampa clyton* tawny emperor ASTCLY

*Battus philenor* pipevine BATPHI WICR

*Cercyonis pegala* common wood nymph CERPEG PIPE WICR HOME

*Erynnis horatius* Horace's dusky wing ERYHOR WICR

*Euphyes vestris* dun skipper EUPVES PIPE WICR

*Staphylus hayhurstii* southern scalloped

> sootywing STAHAY

*Lethe portlandia* pearl eye LETPOR WICR

*Limenitis arthemis* viceroy LIMART WICR

*Megisto cymela* wood satyr MEGCYM WICR EFMO

*Papilio glaucus* tiger swallowtail PAPGLA PIPE WICR HOME EFMO

*Papilo cresphontes* giant swallowtail PAPCRE WICR EFMO

Table 5. Continued.

[Species Common name Taxon code Parks where present]{.underline}
**Woodland species (continued)**

*Papilio troilus* spicebush swallowtail PAPTRO WICR

*Poanes hobomok* hobomok skipper POAHOB WICR

*Poanes zabulon* zabulon skipper POAZAB WICR

*Polygonia comma* comma POLCOM WICR EFMO

*Polygonia interrogationis* question mark POLINT WICR EFMO

*Polygonia progne* gray comma POLPRO EFMO

*Satyrium calanus* banded hairstreak SATCAL EFMO

*Satyrium liparops* striped hairstreak SATLIP EFMO

**Wetland specialist species**

*Calephelis muticum* metalmark CALMUT WICR

*Euphydryas phaeton* Baltimore checkerspot EUPPHA WICR

*Lethe eurydice* northern pearl eye LETEUR PIPE

*Polites mystic* long dash POLMYS PIPE

**Savanna specialist species**

*Callophrys gryneus* olive hairstreak CALGRY WICR EFMO

*Calycopis cecrops* red banded hairstreak CALCEC WICR

*Epargyreus clarus* hoary edge EPACLA WICR EFMO

*Lethe anthedon* wild indigo duskywing LETANT EFMO

*Libytheana carinenta* snout butterfly LIBCAR WICR

*Thorybes pylades* northern cloudywing THOPYL WICR EFMO

*Satyrium edwardsii* Edward's hairstreak SATEDW EFMO

[Table 6. An example of an Access database and its tables and linkages
for storage and handling of the butterfly monitoring data.]{.underline}

**[Tbl_Butterfly_Species]{.underline} [Tbl_Butterfly_Data]{.underline}
[Tbl_Site]{.underline} [Tbl_Weather]{.underline}**

**Species_Code [1]{.underline} Record_ID** \_\_**1** **Site_ID**
1\_\_\_\_\_\_\_∞ Site_ID

TSN  Site_ID **[∞]{.underline}**  Sitecode Event_ID
[**∞**]{.underline} \_\_\_\_\_\_\_\_\_

ITIS data order review Event_ID [**∞**]{.underline} \_\_\_ Unit_Name
Start_Temp 

Genus_and_Species  [∞]{.underline} Species Code  Community_Type
Start_Wind_Speed 

Common_Name Abundance  Park_Code Start_Cloud_Cover 

Guild_Code [∞]{.underline}\_\_\_\_\_\_\_\_\_\_\_\_\_  Date Transect
End_Temp 

EFMO   Datum End_Wind_Speed 

HOME   Coordinate_System End_Cloud_Cover 

PIPE   Zone 

WICR   Northing 

>   Easting 
>
>   Error 
>
>   
>
>   **[Tbl_Event]{.underline}** 

**[Tbl_Butterfly_Guilds]{.underline}**   [1]{.underline} **Event_ID**
1 \_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_\_

**Guild_Code** 1\_\_\_\_\_\_\_\_\_\_\_ Eventcode

Butterfly Guild Description Date

> Collector 1
>
> Collector 2

Figure 1. Number of butterfly species recorded vs. number of transects
sampled for 6 prairies. Prairie designations, followed by prairie area
in ha are:![](media/image3.png){width="9.389583333333333in"
height="4.227777777777778in"}

1\. Tallgrass Prairie National Preserve (12120)

2\. Konza prairie (3473)

3\. Homestead National Monument of America, schoolhouse prairie (0.2)

4\. Warner Park prairie 1 (12.1)

5\. Warner Park prairie 2 (0.8)

6\. Washington prairie (16.2)

APPENDIX A. Locations of butterfly monitoring transects in three parks
of the Prairie Cluster Program. Transect locations for Effigy Mounds
National Monument are not available. 999 indicates no data.

<table>
<colgroup>
<col style="width: 3%" />
<col style="width: 8%" />
<col style="width: 11%" />
<col style="width: 14%" />
<col style="width: 5%" />
<col style="width: 7%" />
<col style="width: 7%" />
<col style="width: 12%" />
<col style="width: 4%" />
<col style="width: 7%" />
<col style="width: 6%" />
<col style="width: 4%" />
<col style="width: 6%" />
</colgroup>
<tbody>
<tr class="odd">
<td><p>Tran</p>
<p>ID</p></td>
<td>TranCode</td>
<td>Unit Name</td>
<td>Community Type</td>
<td>Park</td>
<td>Established</td>
<td>Datum</td>
<td>Coordinate System</td>
<td>Zone</td>
<td>Northing</td>
<td>Easting</td>
<td>Error</td>
<td>Elevation</td>
</tr>
<tr class="even">
<td>1</td>
<td>BH1</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106351</td>
<td>463446</td>
<td>0</td>
<td>365</td>
</tr>
<tr class="odd">
<td>2</td>
<td>BH2</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106338</td>
<td>463468</td>
<td>0</td>
<td>361</td>
</tr>
<tr class="even">
<td>3</td>
<td>BH3</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106366</td>
<td>463461</td>
<td>0</td>
<td>363</td>
</tr>
<tr class="odd">
<td>4</td>
<td>BH4</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106436</td>
<td>463483</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>5</td>
<td>bh5</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106429</td>
<td>463453</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>6</td>
<td>BH6</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106417</td>
<td>463436</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>7</td>
<td>BH7</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>8</td>
<td>BH8</td>
<td>BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>9</td>
<td>NBH1</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>10</td>
<td>NBH2</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>11</td>
<td>NBH3</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>12</td>
<td>NBH4</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106910</td>
<td>463591</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>13</td>
<td>NBH5</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>14</td>
<td>NBH6</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/29</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>15S</td>
<td>4106882</td>
<td>463571</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>15</td>
<td>NBH7</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/30</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>16</td>
<td>NBH8</td>
<td>NORTH BLOODY HILL</td>
<td>GLADE</td>
<td>WICR</td>
<td>05/30</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>17</td>
<td>HOME1</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4461392</td>
<td>683963</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>18</td>
<td>HOME2</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4461424</td>
<td>684070</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>19</td>
<td>HOME3</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/09</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4461402</td>
<td>684112</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>20</td>
<td>HOME4</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/09</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>21</td>
<td>HOME5</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4461561</td>
<td>684154</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>22</td>
<td>HOME6</td>
<td>HOMESTEAD</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4461751</td>
<td>684146</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>23</td>
<td>PIPE1</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/26</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4876913</td>
<td>714494</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>24</td>
<td>PIPE2</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/26</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4877003</td>
<td>714551</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>25</td>
<td>PIPE3</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/26</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4876879</td>
<td>714546</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>26</td>
<td>PIPE4</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/26</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4876879</td>
<td>714623</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>27</td>
<td>PIPE5</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4876484</td>
<td>714429</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>28</td>
<td>PIPE6</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4876297</td>
<td>714399</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>29</td>
<td>PIPE7</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/27</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>30</td>
<td>PIPE8</td>
<td>PIPESTONE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>06/27</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>32</td>
<td><p>PIPE</p>
<p>ORCH1</p></td>
<td>PIPE UNIT 2 WITH ORCHIDS</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>08/14</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>33</td>
<td><p>PIPE</p>
<p>ORCH2</p></td>
<td>PIPE UNIT 2 WITH ORCHIDS</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>08/14</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>34</td>
<td><p>PIPE</p>
<p>ORCH3</p></td>
<td>PIPE UNIT 2 WITH ORCHIDS</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>08/14</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>35</td>
<td><p>PIPE</p>
<p>REST1</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>36</td>
<td><p>PIPE</p>
<p>REST2</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>08/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>37</td>
<td><p>PIPE</p>
<p>REST10A</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>38</td>
<td><p>PIPE</p>
<p>REST10B</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>39</td>
<td><p>PIPE</p>
<p>REST9A</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>40</td>
<td><p>PIPE</p>
<p>REST9B</p></td>
<td>PIPE RESTORED PRAIRIE</td>
<td>TALLGRASS PRAIRIE</td>
<td>PIPE</td>
<td>07/01</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>41</td>
<td><p>PIPE</p>
<p>SOUX1</p></td>
<td>PIPE SOUX QUARTZITE</td>
<td>GLADE</td>
<td>PIPE</td>
<td>08/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>42</td>
<td><p>PIPE</p>
<p>SOUX2</p></td>
<td>PIPE SOUX QUARTZITE</td>
<td>GLADE</td>
<td>PIPE</td>
<td>08/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>43</td>
<td>RAY1</td>
<td>WICR RAY HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>WICR</td>
<td>07/18</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>44</td>
<td>RAY2</td>
<td>WICR RAY HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>WICR</td>
<td>07/18</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>45</td>
<td>SCHOOL1</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462271</td>
<td>683481</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>46</td>
<td>SCHOOL2</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462269</td>
<td>683475</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>47</td>
<td>SCHOOL3</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462267</td>
<td>683468</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>48</td>
<td>SCHOOL4</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462268</td>
<td>683460</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>49</td>
<td>SCHOOL5</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462270</td>
<td>683453</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>50</td>
<td>SCHOOL6</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/10</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td>14T</td>
<td>4462268</td>
<td>683448</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>51</td>
<td>SCHOOL7</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/12</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>52</td>
<td>SCHOOL8</td>
<td>HOME SCHOOL HOUSE</td>
<td>TALLGRASS PRAIRIE</td>
<td>HOME</td>
<td>06/12</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>53</td>
<td><p>WICR</p>
<p>SASV1</p></td>
<td><p>WICR UPLAND SAVANNA/</p>
<p>WOODLAND</p></td>
<td>SAVANNA WOODLAND</td>
<td>WICR</td>
<td>07/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>54</td>
<td><p>WICR</p>
<p>SASV2</p></td>
<td><p>WICR UPLAND SAVANNA/</p>
<p>WOODLAND</p></td>
<td>SAVANNA WOODLAND</td>
<td>WICR</td>
<td>07/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="odd">
<td>55</td>
<td><p>WICR</p>
<p>SASV3</p></td>
<td><p>WICR UPLAND SAVANNA/</p>
<p>WOODLAND</p></td>
<td>SAVANNA WOODLAND</td>
<td>WICR</td>
<td>07/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
<tr class="even">
<td>56</td>
<td><p>WICR</p>
<p>SASV4</p></td>
<td><p>WICR UPLAND SAVANNA/</p>
<p>WOODLAND</p></td>
<td>SAVANNA WOODLAND</td>
<td>WICR</td>
<td>07/13</td>
<td>NAS‑C</td>
<td>UTM/UPS</td>
<td></td>
<td>999</td>
<td>999</td>
<td>0</td>
<td>0</td>
</tr>
</tbody>
</table>

APPENDIX B. Annual report on butterfly monitoring at Homestead National
Monument of America.

**Prairie Cluster
Long-Term**![](media/image1.png){width="6.539583333333334in"
height="1.0743055555555556in"}

**Ecological Monitoring Program**

**Program Report**

**00-001**

**Status Report:**

**1997-1998 Butterfly Monitoring at**

**Homestead National Monument of America**

![](media/image2.png){width="2.1222222222222222in"
height="2.6326388888888888in"}

Status Report:

1997-1998 Butterfly Monitoring at

Homestead National Monument of America

by

William M. Rizzo

U.S. Geological Survey

Northern Prairie Wildlife Research Center

Missouri Project Office

302 Gentry Hall

University of Missouri-Columbia

Columbia, Missouri 65211

August 2000

**1.0 INTRODUCTION**

1.1 [Background]{.underline}

Only about 1% of the original tallgrass prairie ecosystem of the U.S.
still remains (Swengel 1996). The drastic loss of prairie habitat, the
disruption of natural disturbance regimes and nutrient cycles, and the
isolation of the remaining tallgrass prairie habitat make the
preservation of prairie-dependent species a unique challenge (Johnson
and Simberloff 1974; Leach and Givnish 1996; Collins et al. 1998; Kaiser
1998; Schlicht and Orwig 1998). The butterfly communities of tallgrass
prairie remnants are indicators of impacts on a broader spectrum of
invertebrate species (New et al. 1995). Monitoring butterfly communities
offers several advantages: 1) Their habitat preferences and host plant
associations are relatively well-known, which allows classification of
individual species into guilds (Sharp et al. 1974; Scott 1986; Opler and
Malikul 1992; Panzer et al. 1995; Swengel 1996; 1998); 2) A large
literature exists on metapopulation dynamics, dispersal, and the effects
of different habitats on butterfly movement behavior (Sharp et al. 1974;
Fahrig and Paloheimo 1987; Hanski et al. 1995; Hanski et al. 1996; Hill
et al. 1996; Neve et al. 1996; Peterson 1997; Haddad 1999); and 3)
Obligate prairie species respond quickly to changes in native vegetation
(Miller and Harris 1977; Saunders et al. 1991).

In 1997 and 1998, a butterfly species inventory was carried out in
Homestead National Monument of America (HOME) in Nebraska (Mahady and
Debinski 1999). This inventory supplied the data for this report. Two
habitat types, restored prairie and schoolhouse native prairie, were
sampled at HOME (Mahady and Debinski 1999). In 1997, 3 transects in the
restored prairie were sampled in June and 6 in July, and 6 transects
were sampled in June and July of 1998. In the schoolhouse native
prairie, 6 transects were sampled in June and July of 1997, and 8
transects were sampled in June and July of 1998.

1.2 [Objectives]{.underline}

The objectives of butterfly monitoring at HOME are: 1) to determine
changes in the abundance and species richness of each butterfly species
guild by site, and 2) to compare abundance and species richness of the
butterfly species guilds between native and restored prairie areas.

**2.0 METHODS**

2.1 [Field procedures]{.underline}

All butterfly monitoring procedures are described in detail in Debinski
et al. (2000). Transects (5-m wide by 50-m long) were surveyed by a
single observer walking down the middle of the transect at a constant
pace of 10 m per minute. Most butterflies were identified to species in
flight. Individual butterflies that could not be identified in flight
were captured using a butterfly net and then identified. Each transect
was surveyed 6 times during a sampling visit. A minimum of 15 minutes
elapsed between transect counts to allow the butterfly community to
recover from the disturbance of the previous sample. Abundance of each
species that occurred on a transect was recorded on a data sheet.

2.2 [Abundance and species richness of prairie obligate
species]{.underline}

Using the count data sheets, each butterfly species were assigned to a
species guild (Table 1), and the abundance of individuals within each
guild was calculated for each transect count. Species richness was then
calculated as the sum of the total number of species within each guild
for each transect count. After testing for compliance with the
assumptions of analysis of variance, the abundance and species richness
data were analyzed between years by analysis of variance for each
species guild and site using SAS system statistical analysis software
(SAS Institute Inc. 1989-1996). However, because only 2 woodland guild
species and 1 prairie obligate guild species were recorded at HOME
(Table 1), these groups were excluded from analyses of species richness,
because little biological meaning could be ascribed to the findings. In
addition to the interannual comparisons, analysis of variance also was
used on the total dataset to compare abundance and species richness
between the restored and native prairie for each butterfly guild.
Jaccards similarity index also was calculated for further comparison of
the two habitats. The index is calculated as:

SC~J~ = c/(A + B - c),

where c is the number of species occurring in both habitats, A is the
number of species occurring in habitat 1, and B is the number of species
occurring in habitat 2. The index would have a value of 0 if there were
no species in common, and a value of 1 would indicate that all observed
species occurred at both sites.

**3.0 RESULTS**

Tables 2 and 3 show the abundance of each species found on the restored
prairie and native prairie areas, respectively. Table 4 shows the
results of the analyses of variance between years for each butterfly
guild. At the restored prairie site, both the abundance and species
richness of generalist butterfly species differed significantly
(P[\<]{.underline}0.05) between years. Both variables were much higher
in 1998 than in 1997. However, at the native prairie site the abundance
of the prairie obligate butterfly *Speyeria idalia*, the regal
fritillary, was significantly greater in 1997 than in 1998.

The two prairie sites differed significantly (P[\<]{.underline}0.05) in
the abundance and species richness of generalist guild butterflies
(Table 5). Values for both variables were higher in the native prairie
than in the restored prairie. The Jaccard similarity index value for the
generalist guild was 0.54, indicating that the species composition was
not very similar between the sites. The Jaccard index also indicated
that the two habitats were even more dissimilar in the composition of
grassland specialist butterflies (SC~J~ = 0.38), even though the
richness values were very similar (Table 5).

**4.0 DISCUSSION**

It is difficult to explain why the generalist butterfly guild species
were less common and less diverse in 1997 than in 1998. Sampling effort
was probably not a large factor because 9 transects were sampled in 1997
vs. 12 in 1998, both in excess of the number of transects needed to
encounter nearly all the species present (Debinski et al. 2000). Also,
the number of sampling dates per year (2) was equal for both sites.
Additionally, examination of the field conditions during collection
(Table 6) do not suggest physical factors during sampling as a cause for
the observed interannual differences. While flights of some species are
brief but large (Mahady, personal communication, reported 101 sightings
of the powesheik skipper at Pipestone National Monument on 27 June 1998,
but none on 26 June 1997), most of these species are relatively common
species over the entire sampling window (Richard and Heitzman 1987).
Climatological factors may offer the best explanation for the
differences.

In contrast, the single prairie obligate taxon was much more abundant at
the native prairie site in 1997 despite less sampling in that year. At
that site only 1 sample was taken early in 1997, so the actual
difference in abundance may have been underestimated. After a June
emergence, this species is typically present all summer in prairie
habitats, so again, neither ephemeral flight periods, sampling intensity
differences, nor physical weather conditions seem likely to explain the
observed difference at this site. If climatological factors are the
major factor explaining these results then the different guilds must
respond differently to climate forcing.

The site comparisons indicate that the two prairies support much
different abundances and species richness of the most common generalist
butterfly group. Not only do the two sites differ in the diversity of
generalist species, but the Jaccard index also shows that community
composition of the generalist guild also is not very similar. In
contrast, the grassland species guild has the same low diversity but
even less similar species assemblages. Often the number of species
recorded is a function of habitat area. Mahady (1999) found such
significant relationships for butterflies in some geographic regions and
for some guilds but none for the generalist or grassland + prairie
obligate guilds occurring in the Flint Hills region, which includes
HOME. Nevertheless, if this relationship had been found, it would have
predicted greater species richness for the larger restored prairie. The
results of the analyses of variance and the Jaccard indices seem more
likely to be due to underlying differences in the plant communities.
Mahady and Debinski (1999) noted that although native grass and forb
diversity was generally good at both prairies, the restored prairie was
impacted by invasion of woody species and exotic grasses and did not
provide the abundance of nectar sources found at the smaller native
prairie. Additional study would be required to determine if this is a
function of differences in plant community composition between these
sites.

Although the initial limited inventory sampling at HOME precludes the
possibility of assessing causes and effects for these findings, this
data does demonstrate the utility of using butterfly monitoring to
assess ecosystem health. Only by continued monitoring can a dataset be
assembled that will allow testing for long-term trends. For instance,
Selser (1992) concluded that her prairie sites all appeared to have
unique butterfly assemblages and suggested that the intricacies of the
relationships between butterflies, forbs, and grasses would emerge only
with continued monitoring. Thus, a butterfly monitoring program carried
out in conjunction with plant community-(Buck et al. 2000) and
weather-(Akyuz et al. 2000) monitoring programs can provide a powerful
basis for using techniques of multivariate analysis to assess the causes
and effects of findings.

**5.0 REFERENCES**

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Columbia, MO 65211.

Buck, C., G. D. Willson, L. Thomas, M. DeBacker, and W. M. Rizzo. 2000.
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Geological Survey, Biological Resources Division, Northern Prairie
Wildlife Research Center, Missouri Field Station, 302 Gentry Hall,
University of Missouri-Columbia, Columbia, MO 65211.

Collins, S. L., A. K. Knapp, J. M. Briggs, J. M. Blair, and E. M.
Steinauer. 1998. Modulation of diversity by grazing and mowing in native
tallgrass prairie. Science 280:745-747.

Debinski, D., S. Mahady, W. M. Rizzo, and G. D. Willson. 2000. Butterfly
monitoring protocol for four prairie parks. U.S. Geological Survey,
Biological Resources Division, Northern Prairie Wildlife Research
Center, Missouri Field Station, 302 Gentry Hall, University of
Missouri-Columbia, Columbia, MO 65211.

Fahrig, L. and J. E. Paloheimo. 1987. Interpatch dispersal of the
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Haddad, N. M. 1999. Corridor use predicted from behaviors at habitat
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Hill, J. K., C. D. Thomas, and O. T. Lewis. 1996. Effects of habitat
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65:725-735.

Johnson, M. P. and D. S. Simberloff. 1974. Environmental determinants of
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Kaiser, J. 1998. Ecology - bison prime prairie biodiversity. Science
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Leach, M. K. and T. J. Givnish. 1996. Ecological determinants of species
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Mahady, S. 1999. Conservation of tallgrass prairie butterfly species in
a highly fragmented landscape. M.Sc. thesis. Iowa State University.
Ames, IA.

Mahady, S. and D. Debinski. 1999. Habitat management plan for butterfly
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Missouri Field Station, U. S. Geological Survey. Iowa State University,
Ames, IA.

Miller, R. I. And L. D. Harris. 1977. Isolation and extirpations in
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40:57-83.

Neve, G., L. Mousson, and M. Baguette. 1996. Adult dispersal and
genetic-structure of butterfly populations in a fragmented landscape.
Acta Oecologia 17:621-626.

Opler, P. A. and V. Malikul. 1992. A Field Guide to Eastern Butterflies.
Boston: Houghton Mifflin Co.

Panzer, R., D. Stillwaugh, R. Gnaedinger, and G. Derkovitz. 1995.
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insects of the Chicago region. Natural Areas Journal 15:101-116.

Peterson, M. A. 1997. Host - plant phenology and butterfly dispersal -
causes and consequences of uphill movement. Ecology 78:167-180.

Richard, J. and J. E. Heitzman. 1987. Butterflies and Moths of Missouri.
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SAS Institute Inc. 1989-1996. The SAS System for Windows, version 6.12.
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Saunders, D. A., R. J. Hobbs, and C. R. Margules. 1991. Biological
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Journal of the Iowa Academy Science 105:82-88.

Scott, J. A. 1986. The Butterflies of North America. Stanford: Stanford
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Selser, E. J. 1992. Diversity, abundance, and distribution of butterfly
species in remnant and tallgrass prairie. In: Wickett, R. G., Lewis, P.
D., Woodliffe, A., and Pratt, P. editors. Proceedings of the Thirteenth
North American Prairie Conference: Spirit of the Land, Our Prairie
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Sharp, M. A., D. R. Parks, and P. R. Ehrlich. 1974. Plant resources and
butterfly habitat selection. Ecology 55:870-875.

Swengel, A. B. 1996. Effects of fire and hay management on abundance of
prairie butterflies. Biological Conservation 76:73-85.

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tallgrass prairie and pine-barrens. Biological Conservation 83:77-89.

[Table 1. Butterfly species recorded at Homestead National Monument of
America, and classified by ecological guild.]{.underline}

[Species Common Name Species Common Name]{.underline}

**[Open habitat generalist species]{.underline} [Prairie obligate
species]{.underline}**

*Celastrina argiolus* spring azure *Speyeria idalia* regal fritillary

*Colias eurytheme* alfalfa butterfly

*Colias philodice* clouded sulfur **[Woodland species]{.underline}**

*Danaus plexippus* monarch *Cercyonis pegala* common wood nymph

*Euptoieta claudia* variegated fritillary *Papilio glaucus* tiger
swallowtail

*Everes comyntas* eastern-tailed blue

*Hemiargus isola* Mexican blue

*Pholisora cattulus* common sooty wing

*Phyciodes tharos* pearl crescent

*Pieris rapae* cabbage

*Pyrgus communis* checkered skipper

*Speyeria cybele* great spangled fritillary

*Strymon melinus* grey hairstreak

*Vanessa atalanta* red admiral

*Vanessa cardui* painted lady

*Vanessa virginiensis* American painted lady

**[Grassland specialist species]{.underline}**

*Atrytone logan* Delaware skipper

*Chlosyne gorgone* gorgone checkerspot

*Erynnis martialis* mottled dusky wing

*Eurema lisa* little sulfur

*Lycaena hyllus* bronze copper

*Phoebis sennae* cloudless sulfur

*Pieris protodice* checkered white

*Polites peckius* Peck's skipper

*Polites themistocles* tawny edged skipper

Table 2. Species abundance for all sampled transects at the restored
prairie of Homestead National Monument of America in 1997 and [1998.
Nine transects were sampled in 1997 and 12 were sampled in
1998.]{.underline}

[Species Common Name 1997 1998]{.underline}

*Atrytone logan* Delaware skipper 0 1

*Cercyonis pegala* common wood nymph 6 6

*Colias eurytheme* alfalfa butterfly 2 11

*Danaus plexippus* monarch 2 4

*Everes comyntas* eastern-tailed blue 6 8

*Papilio glaucus* tiger swallowtail 1 1

*Pholisora cattulus* common sooty wing 1 0

*Phyciodes tharos* pearl crescent 2 6

*Polites themistocles* tawny edged skipper 1 2

*Speyeria cybele* great spangled fritillary 3 6

*Speyeria idalia* regal fritillary 4 3

*Erynnis martialis* mottled dusky wing 0 1

*Phoebis sennae* cloudless sulfur 0 1

*Pyrgus communis* checkered skipper 0 1

*Vanessa atalanta* red admiral 0 1

*Vanessa virginiensis* American painted lady 0 1

Table 3. Species abundance for all sampled transects at the schoolhouse
native prairie habitat of Homestead National Monument of America in 1997
and 1998. Six transects were sampled in 1997, and 16 were sampled in
1998.

[Species Common Name 1997 1998]{.underline}

*Atrytone logan* Delaware skipper 0 3

*Cercyonis pegala* common wood nymph 3 6

*Colias eurytheme* alfalfa butterfly 2 13

*Danaus plexippus* monarch 6 9

*Euptoieta claudia* variegated fritillary 5 8

*Everes comyntas* eastern-tailed blue 6 16

*Lycaena hyllus* bronze copper 6 4

*Pholisora cattulus* common sooty wing 2 0

*Phyciodes tharos* pearl crescent 0 8

*Polites themistocles* tawny edged skipper 0 4

*Speyeria idalia* regal fritillary 6 9

*Phoebis sennae* cloudless sulfur 0 3

*Pyrgus communis* checkered skipper 0 3

*Vanessa atalanta* red admiral 0 1

*Pieris rapae* cabbage butterfly 0 7

*Polites peckius* Peck's skipper 0 2

*Chlosyne gorgone* gorgone checkerspot 0 1

*Eurema lisa* little sulfur 0 1

*Pieris protodice* checkered white 0 1

*Strymon melinus* grey hairstreak 0 2

*Vanessa cardui* painted lady 0 1

Table 4. Results of analyses of variance for abundance and species
richness between years for each butterfly guild occurring within
[Homestead National Monument of America.]{.underline}

**Restored prairie** Mean ± Standard deviation

[Variable Guild 1997 1998 FPROB Result]{.underline}

Abundance Prairie obligate 2.5 ± 1.3 1.3 ± 0.6 0.2108 Not significant

> Generalist 3.6 ± 2.7 17.2 ± 11.7 0.0031 Significant
>
> Grassland specialist 1.0 1.8 ± 1.0 0.5340 Not significant
>
> Woodland species 2.4 ± 1.7 2.8 ± 1.2 0.6358 Not significant

Species richness Generalist 1.9 ± 0.8 3.7 ± 1.3 0.0014 Significant

> [Grassland specialist 1.0 1.3 ± 0.5 0.6850 Not
> significant]{.underline}

**Schoolhouse native prairie** Mean ± Standard deviation

[Variable Guild 1997 1998 FPROB Result]{.underline}

Abundance Prairie obligate 7.3 ± 4.3 1.2 ± 0.4 0.0008 Significant

> Generalist 18.3 ± 7.3 32.1 ± 25.1 0.2069 Not significant
>
> Grassland specialist 2.0 ± 1.5 2.7 ± 2.5 0.5382 Not significant
>
> Woodland species 2.3 ± 1.5 1.5 ± 1.2 0.4011 Not significant

Species richness Generalist 4.3 ± 1.0 4.3 ± 1.9 0.9803 Not significant

> [Grassland specialist 1.0 ± 0.0 1.5 ± 0.7 0.1101 Not
> significant]{.underline}

Table 5. Results of analyses of variance comparing abundance and species
richness, by species guild, between sites at Homestead National Monument
of America. The two-year butterfly dataset was lumped for these
analyses. RP = Restored prairie; NP = Schoolhouse native prairie

Mean ± standard deviation

[Variable Guild RP NP FPROB Result]{.underline}

Abundance Prairie obligate 2.0 ± 1.2 3.7 ± 4.0 0.3011 Not significant

> Generalist 11.1 ± 11.1 28.4 ± 22.5 0.0034 Significant
>
> Grassland specialist 1.6 ± 0.9 2.5 ± 2.2 0.3992 Not significant
>
> Woodland species 2.6 ± 1.4 1.8 ± 1.3 0.1799 Not significant

Species Richness Prairie obligate 1.0 ± 0.0 1.1 ± 0.3 0.5079 Not
significant

> Generalist 2.9 ± 1.4 4.3 ± 1.7 0.0057 Significant
>
> Grassland specialist 1.2 ± 0.4 1.3 ± 0.6 0.6849 Not significant
>
> [Woodland species 1.8 ± 1.5 3.0 ± 3.0 0.2473 Not
> significant]{.underline}

Table 6. Physical field conditions during the transect surveys at
Homestead National Monument of America.

**Restored prairie**

[Date Transects Temperature (°C) Wind speed (km h^-1^) Cloud cover
(%)]{.underline}

9 June 1997 3 25.5 12 80

10 June 1997 1, 5 25.5-28.9 2-5 15-60

21 July 1997 1-2, 4-6 28.9-32.2 2-5 15-60

22 July 1997 3 32.2 5 60

12 June 1998 1-3 27.8 20-21 25-30

13 June 1998 4-6 25.5 5 0-5

28 July 1998 1-6 30.1 0 0

**Schoolhouse native prairie**

[Date Transects Temperature (°C) Wind speed (km h^-1^) Cloud cover
(%)]{.underline}

22 July 1997 1-6 33.3 2 15

12 June 1998 1-8 27.2 8 30

28 July 1998 1-8 26.8-27.7 15 0-30