The leafhopper species assemblages associated with native and replanted grasslands in southwest
Montana
by James Alexander Bess
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in
Entomology
Montana State University
© Copyright by James Alexander Bess (1997)
Abstract:
Leafhopper (Insecta: Homoptera: Cicadellidae) species assemblages were examined and compared
between four distinct grassland types in southwestern Montana. Three sample sites (or patches) were
chosen within each of the four grassland types (two native and two replanted), for a total of twelve
sites. Leafhopper specimens were collected in sweepnet samples from each of the twelve sites in 1988
and 1991. The leafhopper assemblages from the twelve patches were compared using Spearman’s
correlation analysis to determine which assemblages were most similar. In addition, cluster analyses,
using Goodman-Kruskal’s Gamma coefficient, were performed to give a pictorial representation of
spatial relationships between the assemblages and to compare with the correlation analyses. Analyses
were performed on each individual years’ data and the combined data. The correlation analysis found
the patches with each type to be closely related to one another, although some also correlated closely
with patches from other types. Most of these between type associations were between the native
grassland sites. Correlations were strongest using the combined data. Cluster analyses produced many
spurious associations using the single years’ data that were unsubstantiated by patterns observed in the
raw data. Cluster analysis of the combined data produced associations similar to those observed, with
the correlation analysis and supported by the raw data. A total of 67 leafhopper taxa were identified
during this study, 54 of which are new to Montana. THE LEAFHOPPER SPECIES ASSEMBLAGES
ASSOCIATED) WITH NATIVE AND REPLANTED
GRASSLANDS IN SOUTHWEST MONTANA
by
James Alexander Bess
A thesis submitted in partial fulfillment
o f the requirements for the degree
of
M aster o f Science
m
Entomology
MONTANA STATE UNIVERSITY-BOZEMAN
Bozeman, Montana
May 1997
APPROVAL
o f a thesis submitted by
James Alexander Bess
This thesis has been read by each member o f the thesis committee and has been
found to be satisfactory regarding content, English usage, format, citations, bibliographic
style, and consistency, and is ready for submission to the College o f Graduate Studies.
Kevin O ’Neill
signature)
Approved for the Department o f Entomology
Greg Johnson
(Signature)
Date
Approved for the College o f Graduate Studies
R. L. Brown
(Signature)
Date
STATEMENT OF PERMISSION TO USE
In presenting this thesis in partial fulfillment o f the requirements for a master’s
degree at M ontana State University-B ozeman, I agree that the Library shall make it
available to borrowers under the rules o f the Library.
I f I have indicated my intention to copyright this thesis by including a copyright
notice page, copying is allowable only for scholarly purposes, consistent with “fair use” as
prescribed in the U. S. Copyright Law. Requests for permission for extended quotation
from or reproduction o f this thesis in whole or in parts may be granted only by the
copyright holder.
iii
ACKNOWLEDGMENTS
I would like to thank my wife Karen for her invaluable assistance in the preparation
o f this document. Thanks also to Dr. Kevin O ’Neill and Dr. William Kemp (both with the
Department o f Entomology, Montana State University) for financial and technical
assistance during my masters program. Ms. Catherine Seibert (Department o f Biology,
M ontana State University) has provided invaluable assistance with voucher specimen
preparation and interpolation into the Montana State University Entomology Collection.
Mr. Jeffrey Holmes (Department o f Entomology, Montana State University) collected and
graciously provided the vegetation data used in this document. Mr. David Wachter
(Department o f Entomology, Montana State University) provided invaluable assistance in
the collection o f many samples used for leafhopper-foodplant associations.
I would also like to thank Dr. Andy Hamilton (with Agriculture Canada) and Dr.
Ron Panzer (with Northeastern Illinois University) for their assistance in the determination
o f leafhopper specimens and leafhopper-host plant associations. Dr. Panzer also provided
many useful editorial suggestions during my thesis preparation. Dr. Matthew Lavin
(Department o f Biology, Montana State University) and Dr. Noel Pavlovic (with the U.S.
Geologic Survey at Indiana Dunes National Lakeshore) provided assistance in the
preparation and interpretation o f the statistical analyses used in this study. Finally I would
like to thank Dr. Matthew Ayers and Dr, Roger Strand (at Dartmouth University) for their
assistance in obtaining many o f the references on leafhopper-habitat associations and their
insightful comments during the preparation o f this thesis.
TABLE OF CONTENTS
ACKNOW LEDGM ENTS......................................................................................................... iv
LIST OF TA B LES...........................................
vii
LIST OF FIGURES....................................................................................................................viii
ABSTRACT.................................................................................................................................. ix
INTRODUCTION......................................................................................................................... I
Objectives..................................................................................... ,...................................... 5
H ypotheses........................................................................................................................... 5
MATERIALS AND M ETH O D S....................
7
Selection o f Patches.............................................................................................................7
Sampling Techniques.......................................................................................................... 9
Vegetation Sampling................................................................................................. 9
Leafhopper Sampling............................................................................................... 10
Specimen Identification..................................................................................................... 10
Vegetation........................... :................................................................................... 10
Leafhoppers.............................................................................................................. 11
Analysis............................................................................................................................... 14
Correlation Analysis................................................................................................ 14
Cluster Analysis........................................................................................................ 15
Descriptions o f the Plant Assemblages and Patches..................................................... 16
The Stipa comata/Bouteloua gracilis Association................................................16
The Festuca idahoensis/Agropyron spicatum Association..................................20
The Agropvron cristatum/Mfedicago sativa Association..................................... 22
The Bromus inermis/Medicago sativa Association............................................. 24
R ESU LTS................................................................................................................................... 26
Leafhopper Species Collected........................................................................................ 26
The Stipa comata/Bouteloua gracilis Association......................................................... 26
The 1988 D a ta ......................................................................................................... 26
The Combined D ata................................................................................................. 32
The Festuca idahoensis/Agropyron spicatum Association........................................... 34
The 1988 D a ta ......................................................................................................... 34
The 1991 D a ta ......................................................................................................... 34
The Combined D ata.........................................................................................
35
The Agropyron cristatum/Medicago sativa Association........................................ .....35
The 1988 D a ta ......................................................................................................... 35
The 1991 D a ta ......................................................................................................... 37
The Combined D ata.................
37
The Bromus inermis/Medicago sativa Association....................................................... 37
The 1988 D a ta ......................................................................................................... 37
The 1991 D a ta ......................................................................................................... 39
The Combined D ata................................................................................................. 39
RESULTS: A COMPARISON OF LEAFHOPPER ASSEMBLAGES IN
DIFFERENT PA TCH ES............ ......................................................................................
42
The 1988 D a ta ................................................................................................................... 42
The 1991 D a ta ................................................................................................................... 46
The Combined Data for 1988 and 1991......................................................................... 48
DISCUSSION..............................................................................................................................53
Similarity in Leafhopper Assemblages between Sites................................................... 53
Correlation between Leafhopper Species Abundance and Percent Cover of Know
Host P lants........................................................................................................................ 54
Aceratagallia spp...................................................................................................... 54
Amblvsellus grex......................................................................................................56
Athysanus argentarius.......................................................................
56
Chlorotettix unicolor.......................
56
Diplocolenus confieuratus...................................................................................... 57
Doratura stvlata........................
58
Dorvcephalus platyrhynchus...................................................................................58
Endria inim ica.......................................................................................................... 58
Flexamia abbreviatta and F. flexulosa..............................................................
59
Hecalus spp............................................................................................................... 59
Latalus missellus...........................
60
Orocastus labeculus and 0 . perpusilus..................................................................60
Conclusions Concerning Leafhopper/Host Plant Associations.................................... 61
Leafhopper Taxa occurring Primarily in the Native Plant Assemblages................... 61
Leafhopper Taxa occurring Primarily in the Replanted Plant Assemblages..............63
Leafhopper Taxa with Wide Distributions in the Gallatin Valley (but no well-defined
plant assemblage or host plant preferences)......................
65
Notes on the Rarer Leafhopper Species Recorded in this Study.......... .....................66
N otes on Some o f the Pooled Leafhopper G enera....................................................... 69
SAMPLING EFFORT AND ADDITIONS TO THE LEAFHOPPER
FAUNA OF M O N TA N A .......................................................................................................... 71
Sampling Effort.......... ;..................................................................................................... 71
Additions to the Leafhopper Fauna o f M ontana............................................................74
SUM MARY................................................................................................................................. 76
BIBLIOGRAPHY.......................................................................................................................78
APPENDIX A: 1988 AND 1991 LEAFHOPPER PHENOLOGY DATA FOR THE
TWELVE PATCHES.......■............................................................................ 84
APPENDIX B : THE LEAFHOPPER SPECIES COLLECTED
DURING THIS STU D Y ...................
93
vi
------ ------ ------ ------ ------ --
. .
. I-.
L
. ,
I ,
i i. i _ i _ J i
i i
.
: I I
L IST O F TABLES
TABLE I.
Elevation, Precipitation, and Cover Data for the Sample Patches:
Gallatin County, Montana (1991)..................................................................... 8
TABLE 2.
Plant Species Frequency and Percent Cover at the Native
Habitat Patches (1991 Data).................
TABLE 3.
TABLE 4.
17-18
1988 Leafhopper Species Abundance Data from the
Twelve Patches............................
27
1991 Leafhopper Species Abundance Data from the
Twelve Patches...............................
28
TABLE 5.
Leafhopper Abundance Data by Plant assemblage
(1988-1991 combined data)............................................................................. 29
TABLE 6.
Leafhopper Abundance Data for the Twelve Patches
(1988/1991 Combined).................................................................................... 30
TABLE 7.
Leafhopper Species Richness by Subfamily and Plant
assemblage...................................................................................
31
TABLE 8.
Spearman’s Correlations in Leafhopper Species Abundance
between the Twelve Patches (1988 Data).................................................... 43
TABLE 9.
Spearman’s Correlations in Leafhopper Species Abundance
between the Twelve Patches (1991 Data).....................................................47
TABLE 10.
Spearman’s Correlations in Leafhopper Species Abundance
between the Twelve Patches (1988/1991 Combined D ata)......................... 49
TABLE 11.
Correlation Between Leafhopper Species Abundance
and Percent Cover for Eleven Plant Species...........................
TABLE 12.
55
The Leafhopper Fauna of M ontana..................................... ....... ...................75
vii
:n
/
'I'
I
LIST OF FIGURES
FIGURE I.
Leafhopper Species Relative Abundance - STCO/BOGR
Grasslands (1988 - 1991 combined).............................................................. 33
FIGURE 2.
Leafhopper Species Relative Abundance - FEID/AGSP
Grasslands (1988 - 1991 combined)...............................................................36
FIGURE 3.
Leafhopper Species Relative Abundance - Agcr/Mesa
Grasslands (1988 - 1991 combined).............................................................. 38
FIGURE 4.
Leafhopper Species Relative Abundance - Brin/Mesa
Grasslands (1988 - 1991 combined)........................................................... 40
FIGURE 5.
Cluster Analysis o f the 1988 Leafhopper Data
from the Twelve Patches................................................................................. 45
FIGURE 6.
Cluster Analysis o f the 1991 Leafhopper Data
from the Twelve Patches.........................................;...................................... 50
FIGURE 7.
Cluster Analysis o f the Combined Leafhopper Data
from the Twelve Patches................................................................................. 52
V lll
TH E L E A FH O PPE R SPEC IES A SSEM BLAGES ASSOCIA TED W IT H NATIVE
AND REPLA N TED GRASSLANDS IN SO U TH W EST M ONTANA
James Alexander Bess, 1997
ABSTRA CT
Leafhopper (Insecta: Homoptera: Cicadellidae) species assemblages were examined and
compared between four distinct grassland types in southwestern Montana. Three sample
sites (or patches) were chosen within each o f the four grassland types (two native and two
replanted), for a total o f twelve sites. Leafhopper specimens were collected in sweepnet
samples from each o f the twelve sites in 1988 and 1991. The leafhopper assemblages from
the twelve patches were compared using Spearman’s correlation analysis to determine
which assemblages were most similar. In addition, cluster analyses, using GoodmanKruskal’s Gamma coefficient, were performed to give a pictorial representation o f spatial
relationships between the assemblages and to compare with the correlation analyses.
Analyses were performed on each individual years’ data and the combined data. The
correlation analysis found the patches with each type to be closely related to one another,
although some also correlated closely with patches from other types. M ost o f these
between type associations were between the native grassland sites. Correlations were
strongest using the combined data. Cluster analyses produced many spurious associations
using the single years’ data that were unsubstantiated by patterns observed in the raw data.
Cluster analysis of the combined data produced associations similar to those observed, with
the correlation analysis and supported by the raw data. A total o f 67 leafhopper taxa were
identified during this study, 54 o f which are new to Montana.
INTRODUCTION
Kemp et al. (1990) observed that grasshopper (Orthoptera: Acrididae) species in Gallatin
County, M ontana were distributed non-randomly across grassland sites characterized by different
plant assemblages. Despite variation in grasshopper and plant community composition among
patches within plant assemblages, certain species o f grasshoppers tended to be restricted to xeric,
lower elevation sites at the west end o f the valley, while others showed an affinity for more mesic,
higher elevation sites at the east end. A third group o f species was more widely distributed,
displaying relatively high abundance across the entire valley. These results were confirmed and
expanded by Wachter (1995), who also surveyed the grasshopper communities associated with
grasslands in the Bridger and Hyalite Mountain ranges o f Gallatin County. Similar evidence for
uneven distribution o f grasshopper species across habitats has been reported elsewhere in North
America (Alexander and Hilliard, 1969; Blatchley, 1920; Cantrall, 1943; Evans, 1988; Fielding
and Brusven, 1993, 1995; Joem, 1979, 1982; Mulkern, 1967; Otte, 1981,1984).
Although grasshoppers are often abundant (and sometimes economically important)
herbivores in grassland habitats, leafhoppers (Homoptera: Auchenorrhyncha: Cicadellidae) are
commonly considered the dominant herbivorous insects in temperate grassland ecosystems, in
terms o f density o f individuals and diversity o f species (Cherrill and Rushton, 1993; Waloff, 1973;
W aloff and Solomon, 1972). Studies in Europe and North America have shown that leafhopper
species also exhibit preferential habitat selection.
In England and Europe, numerous studies have been conducted to determine the effects o f
variation in environmental parameters on the composition o f grassland leafhopper assemblages
(Brown, et al., 1992; Morris, 1973, 1981a-b,1990a-b; Novotny, 1994a-b, 1995; Waloff, 1980;
WalofF and Solomon, 1972). These studies can be grouped into four categories: I) effects o f land
management practices (i.e. grazing, mowing or fertilizing); 2) effects o f soil pH; 3) effects o f plant
species composition; and 3) effects o f habitat successional age.
In studies o f grazing effects, Brown et al. (1992) found that the composition o f leafhopper
assemblages on sheep-grazed calcareous grasslands was strongly affected by plant architecture, as
dictated by the particular grazing regime being implemented. Morris (1973) also reported grazing
effects on the composition o f leafhopper assemblages in British grasslands, particularly with
reference to the timing o f the grazing event (i.e. spring vs. fall grazing). In addition, ungrazed
grasslands contained a greater number o f leafhopper individuals and species, when compared with
grazed sites. As with Brown et al., Morris believed grassland structure (especially vegetation
height) was a driving force in determining the composition o f leafhopper assemblages.
Morris (1981a-b, 1983), in studies o f the calcareous grasslands o f England, observed that
cutting o f vegetation during the height o f the growing season (i.e. July) had a significant
deleterious effect on many leafhopper species. Mowing in the spring (May) had a less significant
effect o f shorter duration, being confined mainly to species that reached the adult stage in early
summer. The effects o f the July cutting persisted through winter and into spring, especially in
species that overwinter as adults. Progressive declines were observed in many species over the
course o f a three year study (1973-1975), especially on the plots cut in July. Like Brown and
associates (1992), Morris observed a greater number o f leafhoppers on grasslands that were not
intensively managed.
W aloff (1980) and W aloff and Solomon (1972) compared the leafhopper fauna o f acidic
and calcareous grasslands, noting that some leafhopper species were shared between the two
2
habitats, but in differing proportions. Despite these similarities, there were several leafhopper
species reported only from a single grassland type. Morris (1990a-b), in studies o f leafhopper.
colonization in newly sown calcareous grasslands, observed that the leafhopper fauna o f these
early successional grasslands was dominated by several wide ranging, multivoltine habitat
generalists; with very few o f the rarer, univoltine species characteristic o f more mature, semi­
natural grasslands. Morris concluded that habitat structure, habitat age, and proximity to adjacent
natural grasslands were as important as plant species composition in the determination of
leafhopper assemblages. Brown et al. (1992) also noted that the composition o f leafhopper
assemblages varied greatly between early successional and older, more stable grassland
communities. M uch o f this variation was attributed to the absence o f specific food plants in the
early successional habitats. These plants appeared to require old, relatively undisturbed native
grasslands for survival, thus limiting the distribution o f their associated leafhoppers.
Novotny (1994a-b; 1995), in studies on the grasslands o f central Europe, found that the
majority o f leafhopper species occurring in ruderal, early successional habitats tended to have
wider ranges o f host plant use than species occurring in older, more stable environments. He also
observed that host plant specialization became more prevalent with successional age o f the habitat
and that many o f these specialist species showed poor dispersal capabilities. Leafhoppers that
feed on early successional plant species were also found to have greater dispersal capabilities,
wider ranges o f host plant usage, larger geographic distributions and produced a greater number
o f generations per year than those feeding on more competitive, stress-tolerant plants.
Although age, management history and structure o f grassland plant assemblages has been
shown to have a deterministic effect on resident leafhopper assemblages, the distribution o f many
3
species appears to be associated with the distribution o f their host plants (Brown et al., 1992;
Buntin3 1988; Cherrill and Rushton3 1993; Claridge and Wilson, 1978; Gardner and Usher3 1989;
Genung and M ead3 1969; Hicks and Whitcomb, 1996; Nagel, 1979; Novotny3 1994a-b, 1995;
Prestidge and McNeill, 1983; Teraguchi3 1986; WalofFand Thompson3 1980; Whitcomb et al.,
1986, 1987a-b, 1994).
In N orth America, Whitcomb and associates have conducted numerous studies o f the
grassland leafhopper fauna’s o f the Great Plains and Chihuahuan Desert regions (Hicks and
Whitcomb, 1996; Whitcomb et al., 1986, 1987a-b; 1994). Their studies have focused on three o f
the most species-rich grassland genera in North America; Athysanella: Flexamia and
Laevicephalus. Their results suggest that the distribution o f these leafhoppers is closely
associated with the distribution o f their host plants, although many do not occur throughout the
range o f their host. This non-random distribution within the range o f their hosts indicates that
additional factors are limiting their ability to colonize sites. Therefore, specific grassland types,
while they may have plant species in common with other types, often contain unique leafhopper
assemblages adapted to a specific set o f local environmental parameters.
Because o f their role as the dominant insect herbivores in grassland habitats and their
apparent sensitivity to variations in their local environment, leafhoppers are good subjects for
studying the relationship between insect species distribution and plant community composition.
The focus o f the current study was to determine if the leafhopper species occurring in Gallatin
County, M ontana sorted into relatively discrete groupings according to plant assemblage, as was
observed in the local grasshopper fauna (Kemp et al., 1990). In addition, I wanted to determine if
there were significant differences in leafhopper species composition and relative abundance
4
between native and replanted grasslands within the same plant assemblage. As a final objective, I
wanted to update the list o f leafhopper species in Montana previously compiled by Fox (1924).
Objectives
The goal o f my research was to identify the leafhopper (Homoptera: Cicadellidae) species
assemblages associated with four distinct grassland types (following Mueggler and Stewart, 1980)
in Gallatin County, Montana. Specific objectives o f this project were to:
1.
determine if there are distinct leafhopper species assemblages associated with the
four plant assemblages (two native, two replanted);
2.
determine if the abundance o f selected leafhopper species varies with changes in
percent cover o f known or suspected host plants; and
3.
determine which (if any) leafhopper species are restricted in occurrence to native
grassland patches.
Hypotheses
My first hypothesis is that discrete assemblages o f leafhopper species will be associated
with each o f the four plant assemblages. The null hypothesis would be that leafhopper species are
randomly distributed among the plant assemblages.
My second hypothesis is that the distribution o f leafhoppers will be dependent on the
distribution o f their known or suspected host plants. The null hypothesis is that leafhopper
species distribution is unrelated to the distribution o f their host plants.
5
M y third hypothesis is that some o f the leafhopper species recorded will occur only (or most
abundantly) in the native habitat patches within a particular habitat region. The null hypothesis
would be that leafhopper species are evenly distributed between native and replanted grasslands
within the same habitat region.
6
M A TER IA LS AND M ETH O D S
The vegetation data were collected as part o f rangeland grasshopper studies being
conducted by Dr. William P. Kemp and associates at Montana State University. The leafhopper
specimens used in the present study were collected in sweep net samples used in the abovementioned grasshopper research (Kemp et al., 1990). Throughout this thesis, the terms “patch”
and “site” refer to a defined area within which leafhopper and plant data were collected. Each
patch contained a distinct plant assemblage previously determined by Kemp et al. (1990).
Selection of Patches
Two primary plant assemblages were selected for this study, the Stipa comata Trin. &
Rupr./Bouteloua gracilis (H.B.K.) (STCO /BO G R) and Festuca idahoensis Elmer/Agropvron
spicatum (Pursh) (FEID/AGSP) associations o f Mueggler and Stewart (1980). These represent
dry and mesic grassland types, respectively, and occur along an elevation and precipitation
gradient from 1236 m elevation at Three Forks, (annual precipitation 25-3Ocm), to 1750 m at the
base o f the Bridger Mountains (annual precipitation 40-5 Ocm), Gallatin County, Montana
(longitudes l l l o0 0 '-lll° 4 0 ' east-west, latitudes 46°00'-45°45') (Table I). Two replanted
associations were also studied, the Agropvron cristatum (L.VMedicago sativa L. (AgcryMesa)
and Bromus inermis Levss/Medicago sativa L. (B rin M esa) associations. The replanted
A g c rM e sa stands occur within the same elevation and precipitation zone as the native
ST C O /B O G R grasslands and the replanted B rin M e sa stands occur within the same zone as the
native FEID /A G SP grasslands.
7
Table I. Elevation, Precipitation, and C over D ata for the Sample Patches: G allatin Co., M ontana (1991).
Plant
M ean Annual
Assem blage
Elevation (m)
Precipitation (cm )1
% Grass Cover
% Forb Cover
% Bare Ground
STCO/BOGR (all patches)
,200-140«
20-35
54.50 (mean)
11.10 (mean)
16 20 finnan)
7A
-
-
50.96
20.41
23.88
IOA
-
-
50.21
4.36
10.83
16A
-
-
62.37
8.39
13.75
FEID/AGSP (all patches)
21A
25A
26A
1400 - 2300
-
35-50
-
34.00 (mean)
31.33
35.48
35.17
36.90 (mean)
30.61
30.63
49.42
6.70 (mean)
6.78
2.7
10.63
Agcr/Mesa (all patches)
1200-1400
20-35
38.00 (mean)
IKHmem ,
28.00 (mean,
Tb
-
-
31.69
1.57
16.25
16b
-
-
50.96
6.33
35.25
17b
-
-
30.41
0.85
34.03
Brin/Mesa (all patches)
1400 - 2300
35 - 50
57.00 (mean)
8.50 (mean)
32.00 (mean)
21b
-
-
23.53
18.36
25.14
25b
-
-
67.98
1.09
35.25
26b
-
-
79.24
6.02
34.03
1 Information on precipitation in these Plant Assemblages is taken from Mueggler and Stewart, 1980.
Twelve patches (3 in each o f the 4 plant assemblages) were chosen for use in this study.
Patch selection was based on a Detrended Correspondence Analysis (DCA) o f vegetation
characteristics as outlined in Kemp et al. (1990). Three patches within each o f the two native
plant assemblages, (STCOZBOGR) and (FEID/AGSP), were selected because o f their close
similarities (within assemblage) in plant species composition, precipitation, and elevation (i.e. they
had similar coordinates in the DC A). These native plant assemblages were chosen because they
occurred at opposite ends o f a precipitation and elevation gradient for Gallatin County. Three
patches in each o f the two replanted plant assemblages, (AgciVMesa) and (Brin/M esa), were
chosen using the same criteria. For comparisons between native and replanted patches within a
habitat zone, patches were always adjacent. In order to minimize edge effect, sampling was
restricted to the center o f each patch, avoiding the border between adjacent patches. The
minimum size o f any one patch was 10 hectares and no two distinct native patches shared a
common border.
Sam pling Techniques
V egetation Sam pling
The vegetation within each o f the four plant assemblages was sampled during July and
August o f 1991 to coincide with peak standing cover o f plants (Kemp et al., 1990). Forty 0.10
m2 (20 x 50 cm) quadrats were sampled along a randomly selected transect at each o f the twelve
patches. Within each quadrat, percent canopy cover o f each plant species, litter, moss/lichen, and
bare ground were estimated in 5% increments following the methods described by Daubenmire
9
(1959). Subsequent analyses were based on mean cover-estimates o f plant species over all
quadrats in a patch, as was done in previous studies (Kemp et al., 1990).
Leafhopper Sampling
Sweep nets were used to sample leafhopper populations. Samples were collected at each
o f the twelve patches and consisted o f 200, 180 degree sweeps through vegetation with a 15 inch
diameter net, as described by Kemp et al. (1990). In 1988, samples were collected three times per
year in late May/June, late July,.and late August (Appendix A). In 1991, an additional September
collection was included to sample the fall leafhopper fauna (Appendix A). All sweep samples
were collected between 0930 and 1600 h, under sunny skies (<15% cloud cover) and light winds
(<38 kph). Leafhopper samples were placed either in alcohol (the 1988 samples) or air dried (the
1991 samples). Samples were collected in 1988 and 1991, years o f very poor and fairly normal
rainfall conditions, respectively. These years were selected to gather the greatest possible array o f
leafhopper species, as these insects are known to vary in abundance depending on seasonal rainfall
(Waloff, 1973; Whitcomb et al., 1994).
Specimen Identification
Vegetation
Jeffrey Holmes (with USDA-ARS Rangeland Entomology Laboratory at Montana State
University) identified all plant specimens observed during the vegetation studies. Plant specimens
that could not be identified in the field were returned to the lab for analysis and comparison with
voucher specimens in the Montana State University Herbarium. Most o f the early spring
10
ephemeral genera such as Besseya. Ervthronium. Fritillaria. Dodecatheon. and Delphinium are
conspicuously absent from the species list in Table 2. This is a result o f vegetation surveys being
conducted in mid-summer (to gather information during peak standing cover), when most o f these
spring and early summer species had senesced.
One specific taxonomic note is needed here. Gould (1947) considers all o f the native
N orth American grass species previously assigned to Agropvron to be included in Elvmus. I have
retained the name Agropvron here for the sake o f clarity in comparisons between it and the work
o f Kemp and associates (1990). For leafhopper species which are reported to feed on species o f
Elvmus, I have included Agropyron grasses as potential hosts.
Leafhoppers
When possible, all leafhopper specimens were identified to species. A few groups,
however, are difficult to identify beyond genus because o f similarities in adult features or
unresolved taxonomic problems. Members o f the genus Aceratagallia (subfamily Agalliinae) are
nearly impossible to identify on the basis o f females, and males are difficult as well. This genus is
currently undergoing revision by K.G.A. Hamilton o f the Canada Biosystematic Research
Institute. For the purposes o f the present study, leafhoppers in this genus were studied at the
generic level. A subsample o f male specimens were identified as the species A. sanguinolenta and
A. uhleri. Both species are widely distributed across North America (Beirne, 1956; Delong,
1948).
11
The entire subfamily Typhlocybinae is also problematic, because o f close similarities in
external morphologic and genitalic features, especially in females. These leafhoppers are also very
small (usually <3 mm in length), quite fragile and easily destroyed in sweep net samples. Within
this subfamily, the genera Empoasca and Ervthroneura are difficult to identify and were covered
only superficially by Delong (1948) and Beirne (1956). However, Oman (1949) gives excellent
keys to genera for these groups. Despite the difficulty in identifying these leafhoppers, two taxa
were identified to species. M ost male Dikraneura specimens taken in this study were identified as
D. shoshone. a common grassland species. Given the occurrence o f numerous female specimens
in the samples (without accompanying males), these leafhoppers are analyzed as a genus. Another
grassland species, Forcipata loca. is fairly distinctive and was identified to species. All other
specimens o f this subfamily were identified to genus in this study.
In the subfamily Deltocephalinae, females o f the genera Athvsanella. Laevicephalus.
Limotettix (Scleroracus), and Sorhoanus are difficult (or impossible) to identify to species
(Beime, 1956; Blocker and Johnson, 1990; Delong, 1948; Ross and Hamilton, 1972). There are
also some unresolved taxonomic problems in the genera Scleroracus and Sorhoanus (Hamilton
1992, pers. comm.).
In 1988, an attempt was made to identify specimens o f these four genera to species
(Appendix A). The Athvsanella specimens from 1988 belonged to seven species, acuticauda.
attenuata. occidentalis. robustus. sinuata. terebrans and utahna. In 1991, many females were
encountered, often without associated males. Therefore, given the limitations in identifying
females o f this group, specimens were pooled as Athvsanella spp. All Laevicephalus specimens
collected during this study were females in the “sylvestris-group” (Ross and Hamilton, 1972).
12
Given the general lack o f characters for identifying females o f this genus to species, specimens
were analyzed as Laevicephalus spp. The Sorhoanus specimens from this study were primarily
debilis. although flavidus and orientalis were taken in the mid-elevation mesic patches (Appendix
A). The Scleroracus specimens were largely females with few associated males. The male
specimens o f Scleroracus that were collected belong to the species dasidus and krvptus.
All leafhopper species identifications were made by myself and verified by K.G.A.
Hamilton o f the Canadian Biosystematic Research Institute in Ottawa, Ontario, Canada (except
for the genus Athvsanella. see above). Keys and figures given by Ball and Beamer (1940),
Beamer and Tuthill (1934, 1935), Beime (1952a, b; 1956), Blocker and Johnson (1990), Blocker
and Wesley, 1985; Brown (1933), Crowder (1952), Delong (1926, 1935; 1948), Delong and
Davidson (1935); Delong and Sleesman (1929), Oman (1949), Ross and Hamilton (1972),
Whitcomb and Hicks (1988), and Young and Beirne (1957) were used to identify specimens
sampled during this study. The taxonomy used in this paper follows these works and that of
M etcalf (1964).
Voucher specimens o f all leafhopper taxa identified in this study have been deposited in
the entomology collection at Montana State University, my personal collection or in the Canadian
National Collection at Ottawa, Ontario. A complete list o f the leafhopper species (and their
authors) identified during this study is contained in Appendix B.
13
Analysis
Correlation Analysis
To test the hypothesis that different plant assemblages have different leafhopper
assemblages, I compared the relative abundance o f leafhopper species between patches using
Spearman’s rank correlation analysis. Correlation analyses are useful in determining whether the
distribution o f two or more species are in some way related. A positive correlation between two
species implies that when one increases in abundance, so does the other. A negative correlation
implies that when one species increases in abundance, the other decreases. In community level
analysis, with many species being compared between several sites, pairwise comparisons of
variation in abundance can be performed to gather an overall measure o f similarity (or
dissimilarity) between the sample sites.
Spearman’s rank correlation coefficients were computed for comparisons between each o f
the 12 sample patches using MSUSTAT. Spearman’s rank correlation is a non-parametric test
selected because it works well with non-normally distributed data (Ludwig and Reynolds, 1988).
Only correlation coefficients having a probability o f <0.05 (given the sample size used) were
considered significant. Correlation matrices were generated for each o f the two sample years
(1988 and 1991) and for both years combined. Between patch comparisons did not include zerozero matches in the data set (i.e. a leafhopper species had to be present on at least one o f the two
sample sites to be included in the analysis).
To test the hypothesis that the abundance o f certain leafhopper species correlated with the
percent cover o f known or suspected food plants, I compared the total abundance (1988 and 1991
data combined) o f each leafhopper species on each o f the twelve sites with the percent cover o f
14
both host and non-host plant species (using Spearman’s rank correlation analysis). Host plant
preference was determined through a review o f the literature, discussions with leafhopper experts,
and personal observations.
Cluster Analysis
Cluster analyses are a multivariate procedure used to detect patterns (or groupings) in
data. The relationship between data groups (samples) is usually represented in the form o f a
dendrogram. The length o f the dendrogram branches and the width between them is
representative o f the degree o f relatedness between the samples. Thus, the most similar groupings
are closest to one another in the dendrogram. SYSTAT (Systat, Inc., 1992) was used to perform
cluster analysis o f the leafhopper data from each year and both years’ data combined.
The most crucial part o f performing cluster analyses is the selection o f the distance
measure used to determine the degree o f relatedness between samples. By selecting the wrong
distance measure, spurious results can occur which cloud the interpretation o f the analyses. Given
that the leafhopper data were arranged in a rank order matrix, a distance measure was chosen that
is designed for use with this type o f data arrangement. SYSTAT (in its “CLUSTER” package)
computes the Goodman-Kruskal gamma correlation coefficient for use with rank order data sets.
This distance measure was selected for use in generating dendrograms for the 1988, 1991 and
combined data sets.
15
D escriptions o f the P la n t Assemblages and Patches
M ore than 65 taxa o f plants were recorded during the 1991 sampling period (Kemp et al.,
unpub. data). O f these, 58 were selected for analysis (Table 2). Plant taxa were included in the
analysis if they occurred in three or more o f the 120 sampling plots in at least one o f the four plant
assemblages. The FEID/AGSP grasslands had the greatest plant diversity, with 38 species
recorded. The Brin/Mesa grasslands were the least diverse, with 15 species recorded. Plant
species names in the following descriptions are listed in order o f abundance, from highest to
lowest.
The Stipa comata/Bouteloua gracilis Association
The low elevation (1200-1400m) grasslands surveyed in this study occur on alluvial fans
and plains within the Gallatin Valley and are dominated by Stipa comata and Bouteloua gracilis.
The grasslands sampled in this plant assemblage are semi-arid, with annual precipitation o f 20-35
cm (Mueggler and Stewart, 1980). The patches also had a relatively low diversity o f plant species
(29), when compared with the more mesic native grasslands occurring at higher elevations. Other
important grasses and sedges in the STCO/BOGR type include Poa sandbergii. Koeleria cristata,
Carex filifolia. and Agropvron smithii. Total cover by grasses ranged from 50.2% to 62.4%, with
an average o f 54.5% (Table I).
The most common forb species observed in this plant assemblage include Sphaeralcea
coccinea. Melilotus officinalis, various species o f Antennaria and Astragalus. Vicia americana. and
several species o f Aster. Total cover by forbs ranged from 4.4% to 20.4%, with an average of
16
Table 2. Plant Species Frequency and Percent Cover at the Native Habitat Patches (1991 Data).
N A T IV E P L A N T A S S E M B L A G E S A N D S A M P L IN G P A T C H E S
PLAN T
S P E C IE S
Stipa com ata/Bouteloua gracilis T y p e
7A
FRQ •/. CVR
IOA
FRQ % CVR
16A
FRQ % CVR
Stip a com ata
9 7 .5 0
32.55
100.00
3 6 .0 0
100.00
26.90
B outeloua gracilis
83.00
9.98
7 7 .5 0
3.7 0
100.00
20.43
Festuca idahoensis/A.eropyron spicatum T y p e
TOTAL
FRQ
•/.CVR
99.17
84.83
31.82
11.37
P oa sandbergii
6 7 .5 0
1.85
7 7 .5 0
2 .0 0
6 2 .5 0
2.08
« .1 7
1.98
K oeleria cristata
2 8 .0 0
1.13
2 2 .5 0
0.73
8 2 .5 0
7.78
4433
3.21
Sphaeralcea coccinea
2 7 .5 0
0.28
2 5 .0 0
0 .4 5
4 2 .5 0
1.78
31.47
0.84
4.28
C a rexfilifo lia
10.00
0 .4 0
4 5 .0 0
7.53
3 0 .0 0
4.9 0
2833
Kfelilotus officinalis
6 7 .5 0
17.05
5.00
0.05
12.50
0.73
28-U
5.94
A gropyron sm ithii
6 0 .0 0
3.95
15.00
0.15
25.00
C arex sp. I
A stragalus spp.
5 .0 0
0.05
10.00
0 .1 0
12.50
0.7 3
10.00
0.43
137
0.15
0.44
Vicia am ericana
17.50
1.30
A nten n a ria sp. I
A nten n a ria sp. 2
A ste r spp
2.50
10.00
0.03
0.4 0
30.00
15.00
0.15
17.50
0.18
15.00
12.50
11.47
0 .3 0
32.50
4.58
10.83
2 0 .0 0
0 .2 0
7.50
5.83
7.5 0
0.08
15.00
0.15
0 .4 0
5.00
5.0 0
0.15
7.5 0
O puntia sp.
5.0 0
0.23
P hlox hoodii
5.0 0
0.05
Taraxacum officinalis
F estuca octoflora
15.00
0 .6 0
A gro p yro n spicatum
7.5 0
G utierrezia sarothrae
A rtem esia fr ig id a
5.0 0
10.00
0.53
30 00
4 .0 5
4 2 .5 0
1.45
3 7 .5 0
3.05
2.5 0
0.13
2SA
FRQ •/• CVR
2 0 .0 0
5 .0 0
1.43
26A
FRQ % CVR
FRQ I tV CVR
15.00
0 .8 0
21.67
2.09
3 7 .5 0
1.68
26.67
1.04
14.17
1.14
0.83
0.04
0.38
TOTAL
0.49
1.53
0.08
0.16
5.0 0
0.28
167
0.09
0.05
5.00
0.05
1.67
0.02
5 2 .5 0
6.0 0
SSfO
6.36
3.33
0.10
0.2 8
5.00
5.00
4.17
0.20
0.23
0.28
4.17
0.14
2.5 0
0.03
4.17
0.26
7.5 0
0.18
4.17
0.08
333
0.71
2.13
21A
FRQ •/. CVF
57.50
4.85
10.00
0.3 0
5 7 .5 0
8.23
G uara coccinea
7.5 0
0.18
2.50
0.06
15.00
0.6 0
5.0 0
0 .0 5
2 .5 0
0.13
7.50
0.26
L iatris pu n cta ta
7.50
0.53
2.50
0.18
12.50
0.93
3 2 .5 0
2 .8 5
2 .5 0
0.03
15.83
L27
2.50
0.03
17.50
0.28
5.83
0.09
50.00
0.85
3 0 .0 0
0.65
47.50
10.00
1.03
7.50
1.01
0.74
P lantago pata g o n ica
2 .5 0
0.03
B rom us ja p o n ic u s
5 .0 0
0.05
A llysum desertorum
G naphalium viscosum
2 .5 0
T ragopogon dubius
2 .5 0
5 .0 0
0.05
1.67
0.02
0.83
0.01
62.50
1.53
0.13
0.83
0.04
12.50
1.18
0.03
0.83
0.01
2.5 0
0.03
10.00
0.1 0
15.00
0.55
10.00
0.45
11.67
0 J7
7 7 .5 0
12.18
9 2 .5 0
2 1 .9 0
100.00
23.03
90.00
L ycopodium sp.
30.00
7.53
6 0 .0 0
12.28
6 0 .0 0
11.03
50.00
19.04
10.28
P hlox longifolia
35.00
1.83
5 7 .5 0
2.68
4 2 .5 0
1.73
45.00
2.08
Cerastium arvense
30.00
2.05
30.00
3.38
6 5 .0 0
6.25
3.89
L upinus sericeus
20.00
2.43
5 0 .0 0
6.1 3
52.50
11.18
41.67
40.83
A ch illea m illefolium
30.00
1.00
3 2 .5 0
298
3 5 .0 0
0.98
32.50
1.65
K ficrosteris gracilis
40.00
0.70
2 2 .5 0
0.7 8
2 0 .0 0
0.3 0
27.50
0.59
B alsam orhiza sagittata
22.50
8.25
10.00
3 50
4 7 .5 0
19.00
26.67
10.25
F estuca idahoensis
6.58
B rom us tectorum
32.50
0.55
27 50
1.25
10.00
0.10
23.33
0.63
H ackelia spp.
2 5 .0 0
0 .3 6
4 0 .0 0
0 .8 0
5.00
0.28
23.33
0.48
Stip a viridula
37.50
2.30
2 7 .5 0
1.53
21.67
1.28
7.50
0.08
17.50
0.4 0
3 5 .0 0
0.45
20.00
0.31
30.00
0.30
0.19
A renaria serpiphylla
C ollom ia sp.
D anthonia unispicata
P oa bulbosa
2 2 .5 0
2.45
2 2 .5 0
0.2 3
5.0 0
0.05
19.17
3 2 .5 0
1.38
20.00
2.40
17.50
1.26
2 5 .0 0
0 88
15.83
1.11
C om m andra um bellata
32.50
1.80
7.5 0
0.0 8
2.5 0
0.13
14.17
0.67
P oa pratense
15.00
0.45
2.5 0
0.0 3
2 2 .5 0
0.88
13.33
0.45
P olygonum sp.
25.00
0.25
2 .5 0
0.0 3
9.17
0.09
H eterotheca villosa
2.5 0
0.03
5.00
0.5 3
7.50
0 18
5.00
0.25
7.5 0
0.75
5.00
0.50
4.17
0.42
2.5 0
0.25
A renaria capillaris
R o sa sp.
10.00
2.15
Senecio canus
10.00
2.63
D elphinium sp.
2.5 0
0.03
A gro p yro n cristatum
D raba sp.
K fedicago sativa
D escurainia p in n a ta
Brom us inerm is
A gro p yro n interm edium
P hleum pratense
17
5.00
0.05
4.17
0.80
3J3
0.88
2.50
0.03
Table 2 c o n t'd . Plant Species Frequency and Percent Cover at the Replanted Patches (1991 Data).
R E P L A N T E D P L A N T A S S E M B L A G E S AN D S A M P L IN G P A T C H E S
PLA N T
S P E C IE S
Stip a com ata
A g r o p y r o n c r is ta tu m /M e d ic a g o s a tiv a T y p e
7b
16b
17b
FRQ
% CVR
FRQ
% CVR
FRQ
% CVR
FRQ
% CVR
2.5 0
0.03
12.50
0 .6 5
5.00
0.3 8
6.67
0 J5
2 .5 0
0.38
12.50
0.78
5.00
15.00
0 .2 5
5.00
O Jf
0.08
3,33
0.18
15.83
0.44
33J3
12.42
B outeloua gracilis
P oa sandbergii
K o eleh a c h sta ta
10.00
0.5 3
Sphaeralcea coccinea
C a rexfilifo lia
Mfelilotus officinalis
A gro p yro n sm ithii
C arex sp. I
A stragalus spp.
Vicia a m e h ca n a
12.50
0.33
1 0 0.00
37.25
2 .5 0
0.3 8
2 7 .5 0
0.68
2 0 .0 0
0.3 0
5 7 .5 0
4.13
7 .5 0
10.00
B r o m u s in e r m is /M e d ic a g o s a tiv a T y p e
21b
TOTAL
0.3 0
0.5 0
6.67
0.10
23J3
1.67
FRQ •/. CVR
25b
FRQ
•/• CVR
26b
TOTAL
FRQ
% CVR
FRQ
V.CVR
2.5 0
0.13
0.83
0.04
7.5 0
0.18
2J0
0.06
A n te n n a h a sp. I
A n te n n a h a sp. 2
A s te r spp.
Taraxacum officinalis
15.00
0.45
F estuca octoflora
A gro p yro n spicatum
2 5 .0 0
0.55
5.00
8J3
0.18
0.15
5 .0 0
0.05
1,67
0.02
5.0 0
0.1 5
3.50
0.08
G utierrezia sarothrae
O puntia sp.
P hlox hoodii
A h e m e sia fh g id a
G uara coccinea
L iatris punctata
P lantago pata g o n ica
B rom us ja p o n ic u s
A llysum desertorum
0 .5 0
0.05
5.00
0 .0 5
52.50
0.73
15.00
0 .1 5
22.50
0.29
2.5 0
0 .2 5
0,83
0.08
G naphalium viscosum
T ragopogon dubius
Festuca idahoensis
L ycopodium sp.
P hlox longifolia
Cerastium arvense
L upinus sericeus
A ch illea m illefolium
M icro steh s gracilis
2 .5 0
0.83
0.03
0.01
5.00
0.05
7.5 0
0 .1 8
5.0 0
0.50
4.17
0.23
30.00
0 .4 0
2.5 0
0.03
12 JO
0.16
2.5 0
0.0 3
2 .5 0
0.03
1.67
0.02
1,67
2 .5 0
0.03
4.17
0.02
0.05
24.17
0.63
Balsam orhiza sagittata
B rom us tectorum
H ackelia spp.
Stip a v ih d u la
A renaria serpiphylla
5.00
0.05
C ollom ia sp.
2.50
0.03
7 .5 0
0.0 8
5.00
0.05
6 7 .5 0
1.85
12.50
2.68
2 5 .0 0
2.1 5
92.50
17.55
100.00
20.80
D a nthonia unispicata
P oa bulbosa
C om m andra um bellata
P oa pratense
100.00
40.98
45.83
15.27
52.50
5.25
48.33
7.60
99.17
33.33
4.17
J4.96
5.89
0.12
P olygonum sp.
H eterotheca villosa
A renaria capillaris
R o sa sp.
Senecio canus
D elphinium sp.
A gro p yro n cristatum
D raba sp.
M edicago sativa
D escurainia p in n a ta
B rom us inerm is
A gro p yro n interm edium
1 0 0.00
31.13
1 0 0.00
49.13
2 2 .5 0
0.33
7.5 0
0.18
2.5 0
0.03
100.00
29.25
P hleum pratense
18
100.00 36.50
7.50 0.11
2.50 0.06
0.83 0.01
1 0 0.00
5 7 .2 5
9 7 .5 0
26.83
37.50
6.4 3
6 2 .5 0
11.25
7 .5 0
0 .3 0
5.00
0.05
11.1% (Table I). Cover by bare ground ranged from 10.8% to 23.9%, with an average o f 16.2%
(Table I).
The three patches sampled within this plant assemblage varied with respect to plant
species composition, vegetative cover and plant species relative abundance (Table 2). Patch 7A is
small (ca. 100 hectares) and relatively mesic (in the Agropyron smithii phase o f the STCO/BOGR
grassland type). Surrounded by heavily grazed native and replanted grasslands and agricultural
fields. In addition to the defining grasses o f this assemblage, Agropvron smithii and Melilotus
officinalis are also common. O fthe three patches sampled in this plant assemblage, Patch 7 A had
the greatest number o f plant species (22) and the greatest cover by forbs. Forb cover was
predominantly Melilotus officinalis. Percent bare ground at 7A was the highest o f the three
patches; 7A is grazed by horses and wild ungulates.
Patch IOA is located 5 kilometers south o f patch 7A and represents an area o f moderate
plant species richness (16 species recorded). It is part o f a large native grassland complex
covering approximately 1,000 hectares north o f the town o f Logan, separated from 7A by a series
o f high limestone ridges. M ost o f my observations on leafhopper-foodplant assemblages were
made in this grassland complex (including Patch 9A). In protected areas upslope from the
STCO/BOGR plant assemblage, grasslands in the Agropvron spicatum /Agropyron smithii series
occurred with greater representation by forbs and small shrubs such as Artemesia frigida and
Gutierrezia sarothrae. The area from which vegetation and insect samples were taken is rather
dry and low in plant species diversity, when compared to other patches in this STCO/BOGR
complex. Forb cover and bare ground values were the lowest of the three STCO/BOGR patches
sampled. Patch IOA is lightly grazed by a small herd o f horses and various wild ungulates.
19
Patch 16A is located 25 kilometers south o f the town o f Three Forks and is part o f a
livestock grazed native and replanted grassland complex spreading south along the Madison
River. It is the most intensively grazed o f the three patches in this plant assemblage. Two to
three hundred head o f cattle may be present and the vegetation is often cropped low. Plant
species richness was moderate (19 total) and grass cover totaled 62.4%, the highest value o f the
three patches. Cover by forbs and bare ground was also moderate (8.4% and 13.8%,
respectively). Canopy cover o f Bouteloua gracilis and Antennaria spp. (plants not readily grazed
by cattle (Mueggler and Stewart, 1980) was significantly higher in patch 16A, compared with
patches 7A and I OA. There was a corresponding decrease in coverage by Agropvron smithii and
Stipa comata, species favored by cattle. This condition usually indicates overgrazing (Mueggler
and Stewart, 1980). An unusual finding is the high frequency and cover values for Koeleria
cristata, a species usually favored by cattle. It is unknown why this grass did so well at 16A,
despite the heavy grazing pressure. Mueggler and Stewart (1980) reported similar findings for
this grass species elsewhere in southwest Montana.
T he Festuca idahoensis/A gropyrou sm catum Association
As the valley floor rises north and east into the foothills (1400-23 00m) o f the Bridger
Mountains, there is a corresponding increase in annual precipitation (average 35 to 50 cm). The
grasslands sampled in this region had the greatest plant species richness, o f the four types
sampled, with 39 species recorded from the three patches. The sample patches were dominated
by Festuca idahoensis and Agropvron spicatum. Other important grasses in this community type
include Koeleria cristata. Bromus tectorum, Stipa comata. S. viridula. Danthonia unispicata. Poa
20
bulbosa, ARropyron smithii. and Poa pratensis. Total cover by grasses ranged from 3 1.33% to
35.48%, with an average o f 33.99% (Table I).
The most common forbs recorded from this grassland type include species o f
Lycopodium. Phlox longifolia. Cerastium arvense. Lupinus sericeus, Achillea millefolium.
Microsteris eracilis. Balsamorhiza sagitatta. species o f Hackelia. Arenaria serpiphvlla. a species o f
Collomia. Liatris punctata. Commandra umbellata. and a species o f Polygonum. Total cover by
forbs ranged from 30.61% to 49.42%, with an average o f 36,89%(Table I). Cover by bare
ground ranged from 2.70% to 10.63%, with an average o f 6.70%(Table I).
The three patches in this plant community occur relatively close to one another on the
west slope o f the Bridger Mountains, northwest o f Bozeman. All are roughly 100 hectares in size
and surrounded by a complex o f replanted and native grasslands. Upslope from these grasslands
was a band o f pine and spruce forest interspersed with mesic, forb- and grass- dominated
meadows. Downslope was primarily agricultural land, planted to wheat, oats, and alfalfa. Cattle
grazing also occurred (at varying intensities) on the sample patches. Plant species composition
and percent cover values varied widely between the patches, but species dominance remained
relatively consistent.
Patch 21A had the greatest number o f plant species (32 species), but the lowest cover
value for grasses (31.3%). There was also a high frequency o f weedy species such as Allvsum
desertorum. Cerastium arvense. Bromus tectorum. Poa bulbosa. and Poa pratensis. The regular
occurrence o f these species in native plant communities usually indicates a past history of
disturbance (Swink and Wilhelm, 1994). Bare ground covered 6.8% o f the area sampled.
21
Patch 25A had the lowest plant diversity (29 species), but the greatest cover value for
grasses (35.5%) in the FEID/AGSP association. Danthonia unispicata was a significant member
o f the flora at both this and patch 26A. There was also a moderate frequency o f the abovementioned weedy species, but the lowest frequency o f Pda pratensis. Clubmosses (Lycopodium
spp.) were also a significant component o f the flora. This site had the lowest percentage o f bare
ground (2.7%).
Patch 26A had a high plant species richness (31 species) and a high cover value for grasses
(35.2%). This patch was moderately grazed and had a high frequency o f weedy species such as
Achillea millefolium. Bromus iaponicus. Cerastium arvense. and Poa pratensis. It is adjacent to a
very high quality, lightly grazed FEID/AGSP grassland appearing to have a greater abundance o f
native plant species (Bess, pers. obs.). Bare ground accounted for 10.6% o f the area sampled and
was the highest value for the three patches in this type.
The Agropyron cristatum/Medicago sativa Association
These replanted grasslands were very low in plant species richness, with 16 species
recorded. Cover by grasses was also low, with a corresponding increase in cover by bare ground.
The most abundant grass species on all three patches was Agropvron cristatum. Portions o f the
three sampling areas had not been completely stripped o f the original vegetation, or had
experienced recolonization by native grasses. These grasses appeared in the sample plots as
scattered individuals o f Festuca octoflora. Stipa comata. Bouteloua gracilis, and Poa sandbergii.
Total grass cover ranged from 30.4% to 51,0%, with an average o f 40.5% (Table I). Cover by
native grass species was less than 1.0% (mean) for this plant assemblage. Forbs included Vicia
22
americana. Sphaeralcea coccinea. species o f Draba and Astragalus. Total cover by forbs ranged
from 0.9% to 6.3%, with an average o f 2.9% (Table I).
Cover by bare ground ranged from
25.14% to 35.25%, with an average o f 31.47% (Table I).
Patch 7B had the lowest number o f plant species (7) and was dominated by Agropvron
cristatum. with scattered weedy forbs such as Astragalus spp. and Sphaeralcea coccinea.
Melilotus officinalis was also a minor component. This patch also had the lowest cover by bare
ground (25.1%) o f the three patches sampled in this plant assemblage. Grass cover was moderate
(40.0%) and consisted solely o f Agropvron cristatum.
Patch 16B had a relatively dense cover o f Agropyron cristatum (49.1%), with scattered
individuals o f a few native grasses, such as Bouteloua gracilis. Festuca octoflora. Poa sandbergii.
and Stipa comata. Grasses covered 51.0% o f the area sampled, the highest value among the three
patches. Forbs were dominated by the disturbance-tolerant species Sphaeralcea coccinea and
Vicia americana. Bare ground covered 35.3% o f the area sampled, the highest value o f any o f the
patches sampled.
Patch 17B was relatively depauperate in plant species, vegetative cover consisting of
scattered Agropvron cristatum. Bouteloua gracilis. Stipa comata. Vicia americana and
Sphaeralcea coccinea. Cover by grasses was the lowest o f any patch sampled (30.4%); cover by
forbs was also very low (0.9%). Bare ground comprised 34.0% o f the area sampled, second
highest value for this plant assemblage.
23
The Bromus inermis/Medicago sativa Association
This was the least species rich o f the four grassland types sampled, with 15 plant species
recorded in 1991. The most commonly recorded grasses were Bromus inermis. Poa pratensis.
Agropvron intermedium. Poa bulbosa. and Phleum pratense. All but Poa bulbosa are considered
non-native and are assumed to have been brought in by humans (intentionally or unintentionally).
Poa bulbosa is reported as being widely distributed in disturbed areas and mesic meadows
throughout M ontana (Dorn, 1984). Total grass cover ranged from 23.8% to 79.2%, with an
average o f 57.0% (Table I).
The most commonly recorded forbs were Medicago sativa. Alvssum desertorum. and
Microseris gracilis. Other incidental forbs include; Achillea millefolium. Collomia spp.,
Taraxacum officinalis. Arenaria serpiphvlla. and Tragopogon dubius. M ost o f these forb species
are non-native and weedy in nature. Total forb cover ranged from 1.09% to 18.36%, with an
average o f 8.49 (Table I). Bare ground ranged from 0.70% to 11.73%, with an average o f 6.06%
(Table I).
The three patches sampled in this plant assemblage varied greatly with respect to plant
species dominance and percent cover. Patch 2 IB had the lowest plant species richness (8), lowest
grass cover value (23.53%), and greatest cover by alfalfa (17.55%). Bare ground covered
11.73% o f the area sampled, highest value for this plant assemblage. This patch was vegetated
entirely with disturbance tolerant species such as Allysum desertorum. Bromus inermis. Medicago
sativa. and Poa pratensis. The sample plots contained no Agropyron intermedium or Phleum
pratense. both o f which are present on patches 25B and 26B.
24
Patch 25B also had a low number o f plant species (11 species), but a high cover value for
grasses (67.98%). This patch had no alfalfa present in the sampling plots. A total o f five species
o f grasses were present and Agropvron intermedium. Poa bulbosa. and Poa pratensis were co­
dominants with Bromus inermis. I assume that A. intermedium and P. pratense were planted
along with B. inermis. All other plants were recorded in low numbers and represent disturbancetolerant taxa such as Allvssum desertorum, Hackelia spp. and Microseris gracilis.
Patch 26B had the highest cover value for grasses (79.24%) with Poa pratensis. Bromus
inermis and Agropyron intermedium dominating (in that order). Alfalfa was present, consisting o f
widely scattered individuals. Other plants were minor components o f the flora and consisted o f
weedy, disturbance tolerant species.
25
RESULTS
Leafhopper Species Collected
Overall, a total o f 44,429 leafhoppers, representing at least 67 taxa in ten subfamilies,
were recorded in the 1988 and 1991 samples from the twelve patches (Appendix A). These were
reduced to 58 taxa (see Materials and Methods) for use in analysis (Tables 3-7). O f these 58 taxa,
46 were shared between the native and replanted grasslands, 12 were found, only in the native
types and 7 were recorded only in the replanted habitats (Tables 5 and 6). In the 1988 samples, a
total o f 17,098 leafhopper specimens were recorded from the twelve patches (Table 3), while
27,331 specimens were recorded in 1991 (Table 4). In 1991, only 377 (14%) o f the 27, 331
specimens were collected in the additional September sample (Appendix A). The following
sections present the leafhopper abundance data for the four plant assemblages, by each year and
both years combined. Leafhopper genera and species names are listed in descending order o f
abundance.
The Stipa comata/Bouteloua gracilis Association
The 1988 Data
A total o f 4,347 leafhoppers were recorded from this plant assemblage in 1988 (Table 3).
The most commonly recorded leafhopper species were Orocastus perpusillus. members o f the
genera Athysanella, Aceratagallia and Sorhoanus. Mocuellus caprillus. Orocastus labeculus.
Rosenus cruciatus, Auridius helvus. Cuema striata. Stenometopiellus cookei. and Deltocephalus
26
Table 3. 1988 Leafhopper Species Abundance Data from the Twelve Patches.
GRASSLAND PLANT ASSEMBLAGES AND PATCHES'
LEAFHOPPER
S T C O /B O G R
F E ID / A G S P
S P E C IE S
7A
IOA
16A
T o ta l
21 A
25A
26A
O rocastus perpusillus
438
1788
120
2346
i4
10
Aceratagallia spp.
M ocuellus caphllus
Hebecephalus crueiatus
A uridius he Ivus
Sorhoanus spp.
Orocastus labeculus
24 0
33
97
37 0
79
160
53
98
63
53
18
3
40
21 6
156
I
45
60
72
234
49
9
121
129
21
199
Cuenta striata
Athysanella spp.
Deltocephalus valens
Hebecephalus rostratus
C olladonus montanus
Dikraneura spp.
D orycephalus platyrhynchus
Prairiana cinerea
48
10
45
57
305
33
11
15
I
3
23
10
Endria rotunda
Flexamia abbreviata
Hecalus spp.
Psammotettix lividellus
Colladonus gem inatus
Flexamia Jlexulosa
Frigartus frig id u s
Xerophloea viridis
Stenometopielus cookei
11
8
7
20
6
4
16
3
6
2
2
Empoasca sp.
Streptanus conjinis
Prairiana subta
15
177
271
1413
4
15
20
63
31
225
9
31
61 5
25
18
159
9
10
27
1458
2
2
3
2
20
31
8
i
34
i
3
40
13
4
I
41
5
18
40
13
7
12
19
44
4
2
9
15
91
35
11
137
I
2
65
36 0
97
515
30
17
18
65
2
46
22
38
8
11
18
3
12
I
I
36
20
23
24
2
49
9
3
3
407
27
T o ta l
7
15
3
8
B rin / M e sa
1 7b
26b
T o ta l
1568
8
177
1753
, j ■
I
30
185
2 49
39
39
29
29
I
6
3
4
I
98
73
171
95
3
81
20
190
23
36 6
2
8
9
19
9
3
16
9
35
7
I
29
32
3
I
12
6
79
2
10
9
25
123
10
41
22
114
13
14
10
5
15
I
2
I
16
I
I
2
48
48
15
2
15
2
3
I
I
I
I
3
3
114
66
7
187
I
I
39
9
21
69
I
I
20
23
13
15
I
2
2
I
2
4
6
2
I
: I
2
4
2
15
i
I
2
5
7
107
58
1544
1709
58
7
517
43
5
2153
7
2677
28
4
50
176
77
303
I
11
2
I
I
i
5
5
55
I
I
i
11
2
I
I
3
3
I
2
I
591
67 2
5
128
22
102
130
33
7
40
2
2
2
2
I
I
I
I
Idiodonus aurantiaeus
Lonatura sp. (F)
D ikrella sp.
Amblysellus grex
Balclutha punctata
Nesosteles neglectus
I
2
2
I
2
I
16
I
2
18
2
I
2
I
I
72
2
I
123
Latalus missellus
Athysanus argentarius
Elymana circius
A uridius auratus
Forcipata Ioca
Scaphytopius spp.
S P E C IE S T O T A L
25b
7
I
10
Chlorotettix unicolor
Endria inimica
Doratura stylata
M acrosteles quadrilineatus
D iplocolenus conjtguratus
M esamia coloradensis
Neoeolidia tumidifrons
Lim otettix spp.
Ballana veruta
Laevicephalus sp.
Norvelina seminuda
Elymana sp. (F)
Texananus sp p
IN D IV ID U A L T O T A L
21b
3
I
4
i
Commellus sexvitattus
Pinumius areatus
2
16b
3
i
I
I
7b
3
58
3
3
2
2
231
7
A g cr / M esa
T o ta l
27
6
9
72
72
33
33
3
2
3
2
1277
2365
70S
4347
713
479
602
1794
1674
271
50 6
2451
2119
1055
5332
8506
23
24
14
29
22
22
26
33
21
15
16
28
15
18
27
32
' P l a n t A s s e m b la g e s : S T C O / B O G R = S tip a c o m a ta /B o u te lo u a g r a c ilis ty p e ; F E I D / A G S P = F e s tu c a id a h o e n s is /
A g r o p y r o n s p ic a tu m ty p e ; A g c r /M e s a = A g r o p y r o n c r is ta tu m /M e d ic a g o s a tiv a ty p e ; B r in / M e s a = B r o m u s in e r m is /
K d ed ica g o s a tiv a ty p e .
Table 4. 1991 Leafhopper Species Abundance Data from the Twelve Patches.
GRASSLAND PLANT ASSEMBLAGES AND PATCHES 1
LEAFH OPPER
S T C O /B O G R
F E ID ZA G S P
A g c r/M e s a
S P E C IE S
7A
IOA
16A
Tetal
21A
25A
26A
Telal
7b
Aceratagallia spp.
923
190
411
1524
135
121
76
332
898
M ocuellus caprillus
2 42
31
2
275
17
76
93
6
16b
186
17b
36
B rin / M e sa
Total
1120
21b
25b
26b
Total
649
129
185
96 3
2
i
I
25
28
6
Athysanella spp.
197
69
746
1012
70
28
98
129
305
114
548
Orocastus labeculus
112
32 9
149
59 0
26
76
62
164
2
54
2
58
Sorhoanus spp.
90
83
16
189
49 9
48 7
323
1309
26
14
3
43
35
36
132
203
Psammotettix lividellus
18
9
27
54
35
20
18
73
23
56
67
146
24
96
212
332
O rocastus perpusillus
16
484
16
5 16
95
16
i
112
6
I
124
4 26
I
7
Chlorotettix unicolor
8
34
15
123
6
6
4
I
8
11
74
Dikraneura spp.
76
70
6
152
20
a
7
38
105
197
8
12
26
65
61
2
128
5
9
3
17
69
60
18
147
I
6
I
4
5
10
2
22
24
68
11
47
2
4
18
12
16
46
17
7
I
I
9
I
I
I
3
7
2
2
5
9
11
7
38
56
6
6
12
I
I
I
I
I
I
I
I
M acrosteles quadrilineatus
6
Empoasca sp.
5
Flexamia flexulosa
5
4
59
A uridius ordinatus
4
5
2
Colladonus gem inatus
4
Hebecephalus rostratus
4
6
HecaIus spp.
4
3
C uem a striata
3
I
5
9
Stenometopielus cookei
3
28
I
32
Commellus sexvitattus
I
Dorycephalus platyrhynchus
I
7
49
4
4
4
6
4
2
2
85
4
7
45
52
9
13
107
3
26
29
I
I
13
13
5
A uridius he Ivus
3
3
Rosenus erueiatus
3
3
Flexamia abbreviata
2
2
Frigartus frig id u s
i
I
I
I
I
I
I
Deltocephalus valens
I
Lim otettix spp.
I
I
i
Nesosteles neglectus
I
I
3
Endria inimica
14
D oratura stylata
12
A uridius auratus
7
M esamia coloradensis
3
Laevicephalus sp. (F )
i
I
Scaphytopius spp.
8
8
9
I
3
29
43
i
24
37
I
3
5
5
8
WWm
20
1704
4109
5833
26
2655
6262
8943
7
I
7
i
2
Xerophloea viridis
6
6
12
D iplocolenus configuratus
2
5
7
Balclutha punctata
I
I
Forcipata Ioca
I
I
N eocolidia tum idifrons
I
24
2
2
I
I
I
i
6
6
285
122
431
I
10
11
I
Elymana circius
18
18
6
45
51
Latalus missellus
I
I
26
86
112
Paraphlepsius occidentalis
I
I
3
H ardya dentata
I
4
4
i
Amblysellus grex
Athysanus argentarius
IN D IV ID U A L T O T A L
S P E C IE S T O T A L
1652
20
1267
20
1458
18
4377
27
1213
24
953
22
710
26
2876
35
1139
17
654
13
273
ii
2066
20
1045
13
19
165
188
28
67
95
5282
11684
18011
21
22
27
1 P l a n t A s s e m b la g e s : S T C O / B O G R = S tip a c o m a ta /B o u te lo u a g r a c ilis ty p e ; F E I D / A G S P = F e s tu c a id a h o e n s is /A g r o p y r o n s p ic a tu m typ e;
A g c r /M e s a = A g r o p y r o n c r is ta tu m /M e d ic a g o s a liv a ty p e ; B r in / M e s a = B r o m u s in e rm is /M e d ic a g o s a liv a ty p e .
28
Table 5. Leafhopper Abundance by Plant Assemblage (1988/1991 Combined Data),
GRASSLAND PLANT ASSEMBLAGES 1
LEAFHOPPER
S T C O /B O G R
S P E C IE S
Orocastus perpusillus
Aceratagallia spp.
Athysanella spp.
Orocastus labeculus
M ocuellus caprillus
Sorhoanus spp.
Rosenus cruciatus
Flexamia Jlexulosa
Stenometopielus cookei
A uridius he Ivus
Psammotettix lividellus
C uem a striata
Hebecephalus rostratus
Deltoeephalus valens
Dikraneura spp.
Dorycephalus platyrhynchus
M acrosteles quadrilineatus
Commellus sexvitattus
Colladonus gem inatus
A uridius ordinatus
Hecalus spp.
Prairiana cinerea
Chlorotettix unicolor
Colladonus montanus
Empoasca sp.
Flexamia abbreviate
F rigartusfrigidus
Endria rotunda
Pinumius areatus
Prairiana subta
Xerophloea viridis
Lim otettix spp.
Nesosteles neglectus
Streptanus confinis
Endria inimica
Doratura stylata
Elym ana eireius
D iplocolenus conjiguratus
M esamia coloradensis
A uridius auratus
Neocolidia tum idifrons
Laevicephalus sp. (F)
Ballana veruta
Norvelina seminuda
Scaphytopius spp.
Texananus spp.
Latalus missellus
Forcipata Ioca
Paraphlepsius occidentalis
Balclutha punctata
Lonatura sp. (F)
H ardya dentata
Am blysellus grex
D ikrella sp.
Idiodonus aurantiaeus
Elymana sp. (F)
Athysanus argentarius
Cicadula quinquinotata
T O T A L # I N D I V I D U A L S (N )
F E ID Z A G S P
A g c r /M e s a
% of N
n
• /. o f N
n
% of N
2862
3 3%
139
3%
17
0%
1894
22%
603
5 7%
16%
9%
163
189
13%
4%
2578
1419
13%
2%
B iin / M esa
n
% of N
2716
57
10%
6%
113
2%
592
71
8
0%
I
0%
423
5%
1924
42%
83
2%
45 2
2%
63
1%
21
0%
32
196
1%
4%
15
78 9
491
159
2%
100
80
1%
1%
75
70
1%
4%
40
0%
0%
2
0%
3
0%
661
42
15%
698
0%
1%
7
3%
0%
4
0%
67
1%
1%
55
47
1%
1%
29
1%
18
0%
36
7
0%
175
4%
59 7
2%
38
0%
201
1%
4%
28
0%
9
0%
26
0%
156
3%
17
0%
20 2
16
0%
8
0%
I
0%
14
1%
0%
11
0%
60
1%
17
0%
126
0%
11
0%
49
1%
10
0%
34
i%
153
2
0%
31 0
7%
7
0%
16
0%
2
0%
0%
28
20
0%
0%
3
0%
1
112
2%
7542
28%
2%
I
5
0%
95
18
0%
11620
4 4%
123
0%
I
0%
734
3%
2
0%
33
0%
4
0%
10
0%
8
0%
8
0%
7
0%
5
0%
4
0%
3
0%
2
0%
0%
2
I
I
I
I
0%
i%
0%
0%
52
0%
11
0%
28
0%
2
9
0%
I
0%
0%
0%
0%
0%
11
8
0%
7
0%
6
0%
3
0%
2
0%
I
0%
2
0%
2
0%
8
0%
I
0%
I
0%
I
0%
2 42
1%
I
0%
3
0%
I
0%
i
0%
139
1%
860
3%
2
0%
I
8724
3%
52
4670
0%
TOTAL
S u m (n)
3018
7791
2231
1089
613
2882
243
100
82
HO
1625
131
106
90
1011
37
401
39
214
112
249
12
336
8
SI
5
6
3
2
2
31
30
6
I
7655
11720
141
746
10
40
7
7
2
2
10
2
243
4
I
4
0%
18
0%
4
0%
2
2
0%
0%
2
0%
I
0%
4517
135
1%
i
0%
26518
144
18
4
862
2
2
4
135
i
% of N
7%
18%
5%
2%
i%
7%
1%
0%
0%
0%
4%
0%
0%
0%
2%
0%
1%
0%
0%
0%
1%
0%
1%
0%
0%
0%
0%
0%
0%
0%
0%
0%
0%
0%
17%
26%
0%
2%
0%
0%
0%
0%
0%
0%
0%
0%
1%
0%
0%
0%
0%
0%
2%
0%
0%
0%
0%
0%
44429
1 P l a n t A s s e m b la g e s : S T C O / B O G R = S tip a c o m a ta /B o u te lo u a g r a c ilis ty p e ; F E I D / A G S P = F e s tu c a id a h o e n s is /A g r o p y r o n s p ic a tu m ty p e ;
A g c r /M e s a = A g r o p y r o n c ris ta tu m /M e d ic a g o s a liv a ty p e ; B r in / M e s a = B r o m u s in e rm is /M e d ic a g o s a liv a ty p e .
29
Table 6. Leafhopper Abundance Data for the Twelve Patches (1988/1991 Combined),
LEAFHOPPER
SPECIES
Aceratagallia spp.
Orocastus perpusillus
M ocuellus caprillus
Athysanella spp.
Orocastus labeculus
Sorhoanus spp.
Rosenus cruciatus
A uridius he Ivus
C uem a striata
Deltocephalus valens
Psammotettix lividellus
Hebecephalus rostratus
Dikraneura spp.
Colladonus montanus
Chlorotettix unicolor
Dorycephalus platyrhynchus
Hecalus spp.
Flexamia Jlexulosa
M acrosteles quadrilineatus
Colladonus gem inatus
Empoasca sp.
Prairiana einerea
A uridius ordinatus
Endria rotunda
Flexamia abbreviata
Stenometopielus cookei
Frigartus frig id u s
Xerophloea viridis
Commellus sexvitattus
Pinumius areatus
Streptanus confinis
Prairiana subta
Lim otettix spp.
N esosteles neglectus
Endria inimica
Doratura stylata
A uridius auratus
M esamia coloradensis
D iplocolenus conjiguratus
Laevicephalus sp. (F)
Scaphytopius spp.
N eocolidia tum idifrons
Balelutha punctata
Forcipata Ioca
Elymana circius
Ballana veruta
Norvelina seminuda
Texananus spp.
Latalus missellus
Paraphlepsius occidentalis
Idiodonus aurantiaeus
Lonatura sp. (F)
Elymana sp. (F)
H ardya dentata
D ikrella sp.
Amblysellus grex
Athysanus argentarius
Cicadula quinquinotata
T O T A L # IN D IV ID U A L S
T O T A L # S P E C IE S
GRASSLAND PLANT ASSEMBLAGES AND PATCHES'
F E ID / A G S P
A g c r / Mesa
B r i n / Mesa
S T C O /B O G R
7A
IO A
16A
26A
7B
16B
I7 B
21B
25B
26B
223
508
21A
2 14
2SA
W ii
135
253
2311
204
63
2217
137
362
45 4
2272
136
no
26
3
13
I
3
2
133
2
3
52
69
I
66
31 7
I
5
I
I
40 2
402
84
5
18
4
91
6
242
126
1051
100
45
18
142
6
317
62
3
45
3
4
161
458
170
33
85
71
143
204
76
40
730
646
548
45
18
34
16
32
3
41
98
21
63
12
51
11
11
5
3
20
13
15
21
24
37
19
33
13
8
I
8
7
20
6
4
6
6
88
6
8
12
5
6
I
i
4
3
3
4
5
2
39
2
9
3
6
22
99
27
64
170
5
3
119
3
99
2
8
4 27
10
177
2
94
111
46
18
105
295
197
188
100
22
7
6
5
11
9
8
15
21
6
46
80
74
2
19
12
29
4
7
5
11
70
25
I
5
I
7
29
T 03"
5
9
3
12
5
I
47
I
112
65
25
87
17
2
2
22
24
7
45
13
I
I
2
3
3
2
3
76
I
3
I
2
I
I
I
I
12
15
8
8
7
I
I
I
19
I
3
2
I
2
I
2
9
I
53
9
50
32
24
39
4
I
3
I
I
3
7
I
5
I
127
1762
5653
33
3172
8415
33
7
74
i
I
I
461
199
3
I
2
8
I
I
6
i
I
2
2
123
I
15
6
117
48
I
188
3
I
18
2
2
I
6
I
I
2
I
I
16
i
3
I
2
I
2
I
I
76
28
61
75 6
74
I
2929
29
3632
2163
1926
1431
1312
2813
925
779
3164
6340
17014
27
23
28
27
34
25
20
19
19
26
30
1 P l a n t A s s e m b la g e s : S T C O / B O G R = S tip a c o m a ta /B o u te lo u a g r a c ilis ty p e ; F E I D / A G S P = F e s tu c a id a h o e n s is /A g r o p y r o n s p ic a tu m ty p e ;
A g c r /M e s a = A g r o p y r o n c r is ta tu m /M e d ic a g o s a tiv a ty p e ; B r in / M e s a = B r o m u s in e rm is /M e d ic a g o s a liv a ty p e .
30
T a b le 7. L e a fh o p p e r S p e c ie s R ic h n e ss b y S u b fa m ily a n d P la n t A sse m b la g e .
L E A FH O PPE R
SUBFAM ILY
PLA N T A SSEM BLA G E1
STCO/BOGR
FEID/AGSP
Agcr/Mesa
Brin/Mesa
AGALLIINAE
I
I
I
I
TETTIGONIELLINAE
I
I
I
I
GYPONINAE
2
I
I
I
HECALINAE
I
I
I
I
DORYDIINAE
I
I
O
O
DELTOCEPHALINAE
30
35
26
27
NEOCOELIDIINAE
O
I
I
O
BALCLUTfflNAE
I
2
2
2
TYPHLOCYBINAE
2
3
2
3
LEDRINAE
I
O.
I
TOTALS
.1
40
47
35
37
1P l a n t A s s e m b l a g e s : S T C O / B O G R = S t i p a c o m a ta /B o u te lo u a g r a c i l i s ; F E I D / A G S P = F e s t u c a id a h o e n s i s /A g r o p y r o n s p ic a tu m ;
A g c r / M e s a = A g r o p y r o n c r is ta tu m /M e d ic a g o s a l i v a ; B r i n / M e s a = B r o m u s in e r m is /M e d ic a g o s a l i v a .
31
valens. The remaining leafhopper taxa identified from the 1988 samples each represented less
than 1.00% o f the total recorded for this plant assemblage (Table 3). The 1991 Data
A total o f 4,377 leafhoppers representing 28 taxa were recorded from this plant
assemblage in 1991 (Table 4). The most commonly recorded leafhoppers were species o f
Aceratagallia and Athvsanella. Orocastus labeculus. Orocastus perpusillus. Mocuellus caprillus.
the genus Sorhoanus. Flexamia flexulosa. and Psamotettix lividellus. All other leafhopper taxa
each represent less than 1.00% o f the total number recorded for this plant assemblage (Table 4).
The Combined Data
The low elevation native grasslands o f Gallatin County, Montana were relatively low in
leafhopper species richness (40 taxa total). The most abundant leafhopper taxa recorded from
this plant assemblage were Orocastus perpusillus. several members o f the genera Acerataeallia
and Athvsanella. Orocastus labeculus. Mocuellus caprillus. three species in the genus Sorhoanus.
Rosenus cruciatus. Flexamia flexulosa. and Stenometopiellus cookei (Tables 3-6; Figure I).
Several leafhopper species were recorded almost exclusively from the STCO/BOGR plant
assemblage, although they appeared in low numbers (Tables 3-6; Figure I). These include
Colladonus montanus (8 specimens from patch 7A, 1988 only), Dorvcephalus platyrhynchus
(found primarily in the low elevation native grasslands, (28 specimens), but also in the mid­
elevation ones (nine specimens)), Endria rotunda (three specimens from patch 7A, 1988 only),
Flexamia abbreviatta (five specimens from the STCO/BOGR grasslands and a Bouteloua
specialist), Pinumius areatus (two specimens, patch 16A), Prairiana cinerea (ten specimens from
the STCO/BOGR grasslands, two from the Agcr/Mesa patches), and P. subta (three specimens
32
Figure I. Leafhopper Species Abundance - STCO / BOGR Grasslands (1988-1991 Combined)
10000
! I I I
I!I I l a
S p e c ie s N a m e
IlIIjiiiiiIfIjiiIii
from patch 16A). Each o f these seven species represented less than one percent o f the total
number o f leafhoppers recorded from this plant assemblage.
The Festuca idahoensis/Agropyron spicatum Association
The 1988 Data
A total o f 1,794 leafhoppers representing 29 taxa were recorded from this plant
assemblage in 1988 (Table 3). The most common leafhoppers were members o f the genera
Sorhoanus and Aceratagallia. Chlorotettix unicolor. Psamotettix lividellus, Endria inimica. several
species o f Athvsanella, Rosenus cruciatus. Doratura stylata. Dikraneura shoshone. Deltocephalus
valens. Auridius helvus. Macrosteles quadrinotata. Orocastus labeculus. Mocuellus caprillus and
Hebecephalus rostratus. All other leafhopper taxa identified in the 1988 samples each represented
less than 1.00% o f the total number recorded for this plant assemblage (Table 3).
The 1991 Data
A total o f 2,876 leafhoppers representing 36 taxa were recorded from this plant
assemblage in 1991 (Table 4). Those most commonly recorded were species o f Sorhoanus and
Aceratagallia. Orocastus labecullus, Dikraneura shoshone. Macrosteles quadrinotata. Chlorotettix
unicolor. Orocastus perpusillus. the genus Athvsanella. Mocuellus caprillus. Psamotettix lividellus.
the genus Auridius, Colladonus geminatus. Endria inimica. and Doratura stylata. All other
leafhopper taxa each represent less than L 00% o f the total recorded from this plant assemblage
(Table 4).
34
The Combined Data
The mid-elevation native grasslands had the greatest leafhopper species richness (47 taxa
total) o f the four types sampled (Tables 5 and 6). Samples were dominated by members o f the
genera Sorhoanus and Aceratagallia. Chlorotettix unicolor, several species o f Athvsanella.
Dikraneura shoshone. Orocastus labeculus. Psammotettix lividellus, Doratura stylata, Endria
inimica, Macrosteles quadrinotata and Mocuellus caprillus (Tables 3-6; Figure 2).
Several additional leafhopper species were recorded primarily from this plant assemblage,
although in low numbers (Tables 5 and 6; Figure 2). These include Xerophloea viridis (28
specimens in the FEID/AGSP grasslands, compared with only one from the STCO/BOGR patches
and two from the Brin/Mesa patches), members o f the genus Limotettix (subgenus Scleroracus
(20 specimens from the FEID/AGSP patches, nine from Brin/Mesa and one from the
STCO/BOGR association)), Mesamia coloradensis (eight specimens from the FEID/AGSP
patches, two from the Brin/Mesa association), and Neocolidia tumidifrons (six specimens from the
FEID/AGSP patches, one from the Agcr/Mesa).
The Agropyron cristatum/Medicago sativa Association
The 1988 Data
A total o f 2,451 leafhoppers representing 28 taxa were recorded from this plant
assemblage in 1988 (Table 3). Samples contained a few wide-ranging taxa such as members o f
the genus Aceratagallia, Psamotettix lividellus. Dikraneura shoshone. species o f Hecalus, a few
Athvsanella. Cuerna striata, and the genus Sorhoanus. All other leafhopper taxa each represented
less than 1.00% o f the total number recorded from this plant assemblage (Table 3).
35
Figure 2. Leafhopper Species Abundance - FEID / AGSP Grasslands (1988-1991 Combined)
10000
I I sI I
i I i ^ s
I 11
H i M I
M | H
S p e c ie s N a m e
=: §;
H
The 1991 Data
A total o f 2,066 leafhoppers representing 19 taxa were recorded from this plant
assemblage in 1991 (Table 4). Those most commonly recorded were species o f AceratagalUa and
Athvsanella. Psamotettix lividellus. Orocastus labeculus. Species o f Hecalus, the genus Sorhoanus
and Dikraneura shoshone. All other leafhopper taxa each represent less than 1.00% o f the total
recorded from this plant assemblage (Table 4).
The Combined Data
These grasslands had the lowest leafhopper Species richness (35 species total) o f the four
plant assemblages sampled (Table 5). Samples were dominated by common, widespread taxa
such as the genus Aceratagallia. Psammotettix lividellus. a few species o f Athvsanella. Dikraneura
shoshone. species o f Hecalus and the genus Sorhoanus (2.00%). Other leafhoppers recorded
consistently, but in low numbers, include Colladonus geminatus. Cuerna striata. Hebecephalus
rostratus. Lonatura sp., Macrosteles quadrinotata, Orocastus labeculus. and Rosenus cruciatus
(Tables 3-6, Figure 3). A few leafhopper species were recorded exclusively or predominantly
from this plant assemblage (Tables 3-6). These include an unidentified species o f Lonatura (18
female specimens, only at the Brin/Mesa patches), Hardva dentata (four specimens recorded only
in this plant assemblage), and a species o f Dikrella (two specimens, patch 16b).
The Bromus inermis/Medicago sativa Association
The 1988 Data
A total o f 8,506 leafhoppers representing 32 taxa were recorded from this plant
assemblage in 1988 (Table 3). Samples were dominated by Doratura stylata. species of
37
‘
Figure 3. Leafhopper Species A bundance - Agcr / Mesa Grasslands (1988-1991 Combined)
10000
I ;; i I ;
I ; i
I i s i i
S p e c ie s N a m e
Aceratagallia. Endria inimica. Amblvsellus grex. the genus Athvsanella. Psamotettix lividellus.
Diplocolenus configuratus. the genus Sorhoanus. Dikraneura shoshone. Latalus missellus. and
Balclutha punctata. All other leafhopper taxa each represent less than 1.00% o f the total number
recorded from this plant assemblage (Table 3).
The 1991 Data
A total o f 18,011 leafhoppers representing 26 taxa were recorded from this plant
assemblage in 1991 (Table 4). Those most commonly recorded were Doratura stylata, Endria
inimica. the genus Aceratagallia. Diplocolenus configuratus. Dikraneura shoshone. Psamotettix
lividellus and the genus Sorhoanus. All other leafhopper taxa each represent less than 1.00% o f
the total recorded from this plant assemblage (Table 4).
The Combined Data
These mid-elevation replanted grassland patches were relatively low in leafhopper species
richness (37 species total). Samples were dominated by large numbers o f a few widespread taxa,
such as Doratura stylata. Endria inimica. species o f Aceratagallia. Amblvsellus g re x , Diplocolenus
configuratus. Psammotettix lividellus. Dikraneura shoshone. and members o f the genus
Sorhoanus. The Brin/Mesa patches had the largest leafhopper samples recorded (Tables 3-6,
Figure 4).
39
Figure 4. Leafhopper Species A bundance - Brin / Mesa Grasslands (1988-1991 Combined)
100000
10000 - -
e v i
3 I ! I
I I Ii
I M I I
13 Il I Il
S p e c ie s N a m e
Several species were recorded primarily from this plant assemblage (Tables 3-6). These
include Athysanus argentarius (135 from the Brin/Mesa type only), Auridius auratus (33
specimens from the Brin/Mesa, seven in the FEID/AGSP type), Balclutha punctata (139 from the
Brin/Mesa, four from the Agcr/Mesa and one from the FEID/AGSP types), Elvmana circius (123
from the Brin/Mesa type, 19 in the FEID/AGSP patches), and Latalus misselhis (242 from the
Brin/Mesa patches, one from the FEID/AGSP patches).
41
RESULTS:
A COMPARISON OF LEAFHOPPER ASSEMBLAGES
IN DIFFERENT PATCHES
The Spearman’s correlation analyses allow comparison in the overall similarity between
leafhopper assemblages in different patches for each year and for data combined across years. For
each data set, three types o f comparisons will be made: I) between patches characterized by the
same plant assemblage, 2) between patches characterized by different native plant assemblages
(i.e. STCO/BOGR vs. FEID/AGSP), and 3) between patches characterized by native plant
assemblages and replanted ones within the same habitat region (i.e. STCO/BOGR vs. Agcr/Mesa
and FEID/AGSP vs. Brin/Mesa). Other potential comparisons (i.e. the STCO/BOGR vs.
Brin/Mesa, FEID vs. Agcr/Mesa, and Agcr/Mesa vs. Brin/Mesa) were deemed less relevant
ecologically and are not discussed further in this study. Results o f the cluster analysis are also
included in each o f the following three sections.
The 1988 Data
O fthe 39 potential comparisons between the 12 patches, 15 resulted in significant
between-site correlations in the relative abundance o f leafhopper species (Table 8). Eight o f the
significant correlations occurred between sites characterized by the same plant assemblage. The
leafhopper assemblages at all three STCO/BOGR patches were significantly correlated with one
another, suggesting that similarities in the local plant assemblage results in similar resident
leafhopper communities.
42
Table 8. Spearman’s Correlations in Leafhopper Species Abundance Between the Twelve Patches (1988 data).1
PL A N T A SSEM BLA G ES
Patch
7A
IOA
16A
21A
25A
SIr C O /B O G R
7A
IOA
16A
0.60 *** 0.50
28
24
0.49
24
FEID/AGSP
21A
25A
26A
* 0.38
23
* 0.49
24
0.65
14
NS 0.49
* 0.36
23
* 0.49
* 0.41
25
24
* 0.54
* 0.38
14
14
0.73 *** 0.28
25
30
22
0 .2 1
32
26A
7B
16B
17B
21B
2
Agcr/Mesa
7B
16B
17B
NS 0.34
23
* 0.42
25
NS 0 . 2 2
25
NS 0.29
28
NS 0.28
28
0.16
30
NS 0.08
23
* 0 .1 1
28
NS 0 . 0 2
22
NS 0.30
27
NS 0 . 2 2
27
NS 0.03
32
0.50
25
NS 0.32
21B
Irin/Mesa
25B
26B
NS -0.46
31
NS -0.47
32
NS -0.41
0.51
33
** -0.45
NS 0.25
27
34
* -0.54
NS 0 . 2 0
20
25
28
NS 0 . 2 0 NS -0 . 0 2 NS 0.07
27
26
27
NS 0.19 NS -0.17 NS 0.08
26
26
28
NS' -0.04 NS -0 . 1 0 NS -0 . 0 2
32
32
32
* 0.49
* -0 .2 1 NS -0:15
24
28
27
0.41 NS -0.31 NS -0 .1 1
21
23
25
-0.25 NS -0.31
25
27
0.24
22
21
25B
26B
’Significance levels: * = P<0.05; ** = PO.Ol; *** = P O .001; NS = not significant. The number in the lower left hand comer of each block is the
number of leafliopper species compared between patches minus one.
2Plant A ssem blages: S T C O /B O G R = Stipa comata/Bouteloua gracilis ; F E ID /A G S P = Festuca idahoensis/Agropyron spicatum;
A g c r/M e sa = Agropyron cristatum/Medicago sativa; B rin /M esa = Bromus inermis/Medicago sativa.
** -0.35
41
♦* -0.30
40
** -0.27
35
NS 0.15
33
NS 0.18
35
NS 0 . 1 0
36
NS 0 .0 1
37
NS 0 .1 1
*
NS
NS
NS
NS
NS
NS
NS
33
NS -0.14 NS
34
*
NS 0.45
30
0.64 ***
31
The remaining cases o f close similarity in leafhopper communities were primarily between
the tw o native plant assemblages (STCO/BOGR and FEID/AGSP), where 6 o f 9 correlations
were significant. In contrast, there was very little similarity between leafhopper assemblages at
the native STCO/BOGR patches and nearby sites replanted to AgcrZMesa (one o f nine). This
analysis indicates that degree o f similarity in plant species composition is more important than
spatial proximity in determining the leafhopper fauna o f grassland patches in Gallatin County.
In comparisons among leafhopper assemblages from the FEID/AGSP patches, 21A and
25A were significantly correlated, while 26A did not correlate closely with either patch. None o f
the nine comparisons between the FEID/AGSP and Brin/Mesa patches were significant, again
suggesting that replanting with non-native plant species greatly alters the composition o f the
resident leafhopper fauna. Leafhopper samples from patches within each o f the non-native plant
assemblages showed some similarity with each other (2 o f 3 comparisons significant within both
the Agcr/Mesa and Brin/Mesa types), although most o f these significant correlations were
relatively weak.
The cluster analysis performed on the 1988 leafhopper data resulted in an arrangement o f
the 12 patches (Figure 5) different from that obtained with the correlation analyses. One striking
result was the clustering o f several STCO/BOGR patches with ones from the Brin/Mesa
assemblage (i.e. 7A and IOA with 21b; 16A with 26b). In fact, patches 7A (STCO/BOGR) and
21b (Brin/Mesa) were the sites most closely associated in the dendrogram. The correlation
analysis showed 7A and 21b to have a significant, but negative (rs = -0.46, P O . 01) relationship.
44
Figure 5. C luster Analysis of the 1988 L eafhopper Data from the Twelve Patches1
Distance Measure is I minus Goodman-Kruskal's gamma coefficient
Single Linkage Method (Nearest Neighbor).
Plant
Assemblage1 Patch
FEIDZAGSP
26 A .
STCOZBOGR
IOA
STCOZBOGR
7A
BrinZMesa
21b
FEIDZAGSP
21A
FEIDZAGSP
25A
AgcrZMesa
16b
BrinZMesa
25b
BrinZMesa
26b
STCOZB OGR
16A-
AgcrZMesa
7b
AgcrZMesa
17b
DISTANCES
1.000
'Plant Assemblages: STcOZBOGR=Szzpa comata/Bouteloua gracilis; FElDZAGSP=FesZMca idahoensis/Agropyron spicatum ;
AgcrZMesa = Agropyron cnstatumfMedicago sativa; BrinZMesa = Bromus inermis/Medicago sativa.
45
The correlation between IOA and 21b was also negative and significant (rs = -0.47, P<0.01). In
contrast, patches 21A and 25A were also closely associated in the dendrogram, a condition that is
supported by the raw data and correlation analysis. Patch 7b also associated closely with patch
17b, and the two formed an outlier group relative to the rest o f the patches. Site 16b, however,
clustered quite closely with most o f the patches, an association unconfirmed by either the
correlation analysis or the raw data.
The 1991 Data
O fthe 39 potential comparisons between patches, 13 were significant (Table 9). Ten o f
these significant correlations occurred between patches characterized by the same plant
assemblage. As in 1988, the leafhopper assemblages at all three STCO/BOGR patches were
significantly correlated with one another, but showed very little similarity to the nearby non-native
Agcr/Mesa patches (i.e. only I o f 9 comparisons were significant). Other similarities with the
1988 data include: I) leafhopper assemblages at 26A being uncorrelated with those from 21A or
25 A and 2) leafhopper assemblages at the FEE)/AGSP patches being uncorrelated with those
from the Brin/Mesa patches. Leafhopper samples from patches within each o f the two non-native
plant assemblages were more closely correlated in 1991 than 1988 (i.e. all comparisons significant
in both Agcr/Mesa and Brin/Mesa). Perhaps the greatest difference between the 1988 and 1991
leafhopper samples was the reduction in similarity between the two native plant associations (i.e.
only 2 o f 9 correlations were significant, compared with 6 o f 9 in 1988).
f
46
Table 9. Spearm an’s Correlations in Leafhopper Species Abundance Between the Twelve Patches (1991 data).1
PLA N T ASSEM BLA G ES2
Patch
7A
IOA
16A
21A
25A
26A
7B
16B
17B
21B
STCO/BOGR
7A
IOA
16A
FlEID/AGSP
21A
25A
26A
0.65 *** 0.65 *** 0.64 *** 0.58 ** 0.23
24
23
28
27
32
0 . 6 8 *** 0.39
0.28
NS
-0.06
*
24
30
32
33
0.35
N 0.35 NS -0.04
28
28
32
0.57 *** 0.27
31
32
0.28
31
Agcr/Mesa
7B
16B
17B
Brin/Mesa
21B
25B
26B
NS 0.72 *** 0.45
* 0.40 NS 0.03
21
23
22
26
NS 0.48
* 0.40 NS 0.30 NS 0.21
23
23
22
27
N 0.35 NS 0.40 NS 0.31 NS -0.13
23
22
21
24
NS 0.65 *** 0.47 * 0.35 NS 0.39
27
25
25
26
N 0.60 ** 0.52 * 0.45
* 0.30
30
23
24
26
0.22 NS 0.10 NS 0.01 NS 0.28
30
29
29
29
0.57 ** 0.62 ** 0.18
20
19
23
0.80 *** 0.20
14
19
0.09
17
NS -0,21
28
NS -0.28
32
NS -0.30
30
* 0.16
30
NS 0.04
28
NS 0.26
32
NS -0.09
27
NS 0.00
24
NS -0.05
23
0.60
23
N -0.18 N
31
N -0.27
33
NS -0.26
30
NS 0.02
31
NS -0.06
29
NS 0.19
31
NS -0.09
30
NS 0.12
26
NS 0.15
25
** 0.50
24
25B
0 79
25
26B
'Significance levels: * = P O .05; ** = P O .01; *** = PO.OOl; NS = not significant. The number in the lower left hand comer of each
block is the number o f leafhopper species compared between patches minus one.
2P la n t A s s e m b la g e s : S T C O /B O G R = Stipa com ata/Bouteloua g ra c ilis ; F E ID /A G S P = Festuca idahoensis/Agropyron spicatum ;
A g c r/M e s a = A gropyron crisiatum /M eaicago s a tiv a ; B r in /M e s a = Brom us inermis/M edicago sativa.
N
NS
NS
NS
NS
NS
NS
NS
*
***
As observed in 1988, cluster analysis o f the 1991 leafhopper data resulted in some very
unusual associations between the twelve patches, although they were more closely related than in
1988 (Figure 6). Patch 7A was most closely associated with 16b, and this cluster (7A and 16b)
with 16A. Patch 7A was only weakly associated with 16b in the correlation analysis, being most
closely related to IOA both years (Tables 8 and 9). In the cluster analysis, IOA was not closely
associated with 7A, being one o f the least dissimilar o f the twelve patches compared. As in 1988,
Goodman-KruskaV s gamma coefficient analysis may not be approbate for this data set.
The Combined Data for 1988 and 1991
O fthe 39 potential comparisons using the combined leafhopper data from the twelve
patches, 22 were significant (Table 10). All twelve o f the potential comparisons between patches
characterized by the same plant assemblage were significant at P < 0.01 or P < 0.001. Thus,
combining data from two years with different precipitation patterns resulted in an even greater
indication that patches with similar plant species composition tend to have similar leafhopper
assemblages.
Combining results for the two years did not alter the conclusions that leafhopper
assemblages differ between the FEDD/AGSP and Brin/Mesa patches. However, whereas in 1988
and 1991 there was little indication o f significant correlation in the leafhopper assemblages o f the
STCO/BOGR and Agcr/Mesa patches, significant correlations appeared in 6 o f 9 comparisons
using the combined data. Thus, differences observed between the leafhopper assemblages at the
STCO/BOGR and Agcr/Mesa patches become
48
Table 10. Spearman’s Correlations in Leafhopper Species Abundance between the Twelve Patches.
(1988/1991 combined data).1
PLA N T ASSEM BLAG E2
Patch
7A
IOA
16A
21A
25A
26A
7B
16B
17B
21B
25B
STCO/BOGR
7A
10A
16A
F]EID/AGSP
21A
25A
26A
Agcr/Mesa
7B
16B
17B
0.72 *** 0.64 *** 0.42 ** 0.48 ** 0.30 * 0 . 6 6
31
32
38
43
36
34
0.62 *** 0.42 ** 0.34
* 0.17 NS 0.55
30
37
36
43
33
0.32 NS 0.31 NS 0.05 NS 0.38
34
32
42
32
0.74 *** 0.43 ** 0.52
33
38
35
0.49 ** 0.57
40
38
0.38
38
*** 0.19
36
*** 0.26
34
* 0.17
31
** 0.45
35
*** 0.43
32
Brin/Mesa
21B
25B
26B
NS 0.45 ** -0.19
34
37
NS 0.42 ** -0.21
32
36
NS 0.40 * -0.28
27
31
** 0.38 * 0.1
34
32
* 0.43
* 0.21
30
31
* 0.12 NS 0.12 NS 0.15
42
40
37
0.41 ** 0.57 *** 0.04
31
29
33
0.63 *** 0.11
24
28
0.06
27
NS -0.24
40
NS -0.23
39
NS -0.31
37
Ns 0.25
36
NS 0.21
34
NS 0.21
39
NS 0.01
35
NS 0.07
NS -0.21
44
NS -0.23
43
NS -0.21
40
NS 0.15
36
NS 0.17
33
38
NS
NS
NS
NS
NS
37
NS 0.24 NS
40
NS 0.04 NS
40
NS 0.16 NS
NS 0.10 NS 0.02 NS
33
37
0.52 *** 0.48 **
29
33
0.76 ***
33
‘Significance levels: * = P<0.05; ** = P<0.01; *** = P O .OOI; NS = not significant. The number in the lower left hand comer of each block is the
number of leafhopper species compared between patches minus one.
2P la n t A sse m b la g e s : S T C O /B O G R = Stipa com ata/Bouteloua g ra c ilis ; F E ID /A G S P = Festuca idahoensis/Agropyron spicatum ;
A g c r /M e s a = Agropyron cristatum /M edicago sa tiv a ; B r in /M e s a = Brom us inermis/Medicago sativa.
Figure 6. C luster Analysis of the 1991 Leafhopper D ata from the Twelve Patches1
Distance Measure is !minus Goodman-Kruskal s gamma coefficient
Single Linkage Method (Nearest Neighbor).
P la n t
A s s e m b la g e 1
P a tc h
AgcrZMesa
17b
STCOZBOGR
16A
AgcrZMesa
16b
STCOZBOGR
7A -
AgcrZMesa
Tb
BrinZMesa
26b-
BrinZMesa
25b
FEIDZAGSP
25A -
FEIDZAGSP
26 A -
FEIDZAGSP
21A
STCOZBOGR
IOA-
BrinZMesa
21b-
DISTANCES
0.500
lPlant Assemblages: STCOZBOGR=Stipa comata/Bouteloua gracilis; FElDZAGSP=Fejfuca idahoensis/Agropyron spicatum;
AgcifMes^=Agropyron cnstatum/Medicago saliva; BrinZMesa=BroniMJ inermis/Medicago saliva.
50
less evident when compared over a longer time interval. The leafhopper fauna’s o f the
STCO/BOGR and FEDD/AGSP patches remained somewhat closely correlated, with four o f nine
comparisons significant.
In contrast to the dendrograms produced with the individual sampling year’s data, the
cluster analysis o f the combined data resulted in associations that more closely matched the
correlation analyses and patterns observed in the raw data (Figure 7; Tables 6 and 10). The
combined data dendrogram showed patches 7A and IOA to be most closely associated, followed
by 16A (all within the STCO/BOGR plant assemblage). The cluster containing these sites was
then most closely associated with a cluster containing two o f the native mid-elevation patches,
21A and 25 A. These five patches formed a fairly discrete, native grassland cluster.
The next discrete cluster in the dendrogram is composed o f patches 7b, 16b and 17b in the
Agcr/Mesa assemblage. Patch 21b is an outlier associated with the above-mentioned clusters.
The final cluster in the dendrogram (containing patches 26A, 26b and 25b) is very interesting and
reflects patterns observed in the correlation analysis and the raw data. These three sites contained
many rare and unusual leafhopper taxa, resulting (at least in part) from the high diversity of
observed plant species. Although these sites were closely associated with one another in the
dendrogram, they form a distinct outlier group distantly related to patch 21b and the remaining
sample sites.
51
Figure 7. C luster Analysis of the Com bined Leafhopper Data from the Twelve Patches1
Distance Measure is l-Goodman-Kruskal's gamma coefficient.
Single Linkage Method (Nearest Neighbor).
P lan t
A ssem blage1
Patch
Brin/Mesa
21b
AgcrZMesa
17b
AgcrZMesa
16b
AgcrZMesa
Tb
FEIDZAGSP
2 5A -
FEIDZAGSP
2 1A -
STCOZBOGR
16A-
STCOZB OGR
TA -
STCOZBOGR
IOA-
FEIDZAGSP
26 A -
BrinZMesa
2 6b -
BrinZMesa
25b-
DISTANCES
1.000
lPIant Assemblages: STCOfBOGR-Stipa comata/Bouteloua gracilis; FEIDZAGSP=FesfMca idahoensis/Agropyron spicatum ;
AgCTfMssa=Agropyron cnstatum/Xledicago saliva ; BrinZMesa=SromMs inemiis/Medicago saliva.
52
DISCUSSION
Similarity in Leafhopper Assemblages between Sites
Overall, the correlation analyses support the hypothesis that leafhopper assemblages
occurring on grassland patches in Gallatin County tend to be most similar among sites
characterized by the same plant assemblage. The leafhopper assemblages occurring on native
grasslands were also found to be mostly uncorrelated with those on nearby replanted grasslands,
although samples from several o f the STCO/BOGR and Agcr/Mesa patches did correlate
significantly using the combined 1988/1991 data. In addition, many significant correlations
occurred when comparing the leafhopper assemblages from the STCO/BOGR and FEID/AGSP
patches.
The cluster analysis produced many unusual and spurious associations when used on the
individual year’s leafhopper data. However, when the combined data were analyzed, groupings
very similar to those determined by plant assemblage were observed. The STCO/BOGR
grasslands were the most closely associated, followed by two o f the FEID/AGSP patches. The
Agcr/Mesa patches were also found to be closely associated, although patches 26A, 25b and 26b
formed a distinctive outlier, closely associated with one another but not the remaining patches.
This pattern mimics that shown in the plant data, with the native patches being most similar to
onne another (see Table 2). Patches 26A, 25b and 26b also had unusual vegetative components
(i.e. all three having high cover values for Poa pratensef when compared to the remaining sites in
their plant assemblage.
53
Correlation between Leafhopper Species Abundance and Percent Cover of Know Host
Plants
H ost plant associations are unknown for many species o f Cicadellidae. However, several
o f the species collected in this study are known well enough to allow testing o f the hypothesis that
their abundance across the twelve patches correlates with percent cover o f known or suspected
host plants (Table 11). The following is a discussion o f the thirteen leafhopper taxa used in this
comparison and how their abundance correlated with percent cover o f known or suspected hosts.
Aceratasallia spp.
Aceratagallia spp. (primarily A. sanguinolenta and A. uhleri) were relatively abundant
across all patches (Table 6). Their overall abundance correlated highly with percent cover by
plants in the family Fabaceae (Table 11), their known host plants (Beirne, 1956; Delong, 1948).
Aceratagallia sanguinolenta. for example, is a common transcontinental species known as the
clover or alfalfa leafhopper (Beirne, 1956; Delong, 1948). O f the twelve patches sampled,
Aceratagallia leafhoppers were most abundant at sites 7 A, 16A and 7b, all o f which had high
cover values for yellow sweet clover IMelilotus officinalis! and site 21b which had abundant
alfalfa IMedicago sativa). Given the prevalence o f these plant species in disturbed habitats
throughout the county, they perhaps serve as the major local hosts for these leafhoppers.
In addition to occurring in disturbed Fabaceae-dominated sites, Aceratagallia leafhoppers
were observed in a variety o f native plant associations where the only potential hosts were native
Fabaceae. These potential host plants were primarily species o f Astragalus (in low elevation
Stipa/Bouteloua grasslands and in alpine tundra) and Lupinus (in the Festuca
idahoensis/Agropvron spicatum grasslands and other montane meadow communities). Members
54
Table 11. Correlation between Leafhopper Species Abundance and Percent Cover for Eleven Plant Taxa (1991 data)1.
Plant Taxa2
Leafhopper Taxa
Agropyron
spp.
A . spicatum,
Agropyron A . spicatum,
A . smithii,
cristatum
A . sm ithii A . intermedium
Bouteloua
eracilis
Poa
spp.
Acertagallia spp.
NS
NS
NS
NS
NS
NS
Amblysellus grex
NS
NS
-0.75**
NS
NS
NS
Athysanus argentarius
NS
NS
NS
NS
NS
Chlorotettix unicolor
NS
NS
0.68*
0.65*
NS
Diplocolenus configuratus
NS
NS
NS
NS
Doratura stylata
NS
NS
NS
Dorycephalus platyrhynchus
NS
NS
Endria inimica
NS
Stipa
comata
Fabaceae
NS
0.66*
-0.87***
NS
0.69*
-0.58*
NS
NS
NS
NS
-0.85***
0.89***
-0.71**
NS
0.59*
-0.75**
0.92***
-0.61*
NS
0.62*
NS
NS
0.77**
NS
NS
NS
NS
-0.85***
-0.67*
NS
-0.58*
NS
NS
NS
0.85***
NS
0.74**
NS
0.88***
0.89**
NS
NS
NS
NS
NS
NS
Latalus missellus
NS
NS
NS
NS
NS
0.82***
-0.72**
NS
Orocastus labeculus
NS
NS
-0.83*
NS
0.67*
-0.62*
0.94**
NS
NS .
NS
0.64*
NS
0.61*
-0.58*
0.95**
NS
Flexamia Jlexulosa
Hecalus major
Orocastus perpusillus
1Significance levels: * = P O .05; ** = P O .01; *** = PO.OOl.
2 Correlations in BOLD are known or suspected food plants for these leafhopper species.
NS
0.96***
o f these plant genera are common throughout southwestern Montana and likely serve as major
hosts for these leafhoppers in more pristine, native plant assemblages. Over twice as many
Aceratagallia leafhoppers were collected in the replanted patches (Table 6).
Amblvsellus grex
Beime (1956) reported A. grex as being widely distributed in southern British Columbia.
The host plant o f this leafhopper is unknown and its abundance did not correlate positively with
percent cover values for any o f the dominant grass species occurring in the four plant assemblages
(Table 11). The closely related A. curtisii is reported to feed on bluegrass (i.e. Poa spp.) in the
Midwest (Delong, 1948).
Athvsanus argentarius
Athysanus argentarius is a non-native leafhopper species introduced to eastern Canada from
Europe in 1940 (Beime, 1956). From eastern Canada, A. argentarius has spread westward to
Manitoba and Montana, and southward to Iowa and Kentucky (Beirne, 1956; Bess, pers. obs.;
Hamilton, 1983). Athysanus argentarius is reported to feed on a variety o f grasses and its
abundance at the twelve patches correlated strongly with percent cover by Poa spp. (Table 11).
Chlorotettix unicolor
The genus Chlorotettix is associated primarily with grasslands and wetlands in the Great
Plains and Midwest (Blocker and Reed, 1976; Cwikla, 1987; Delong, 1948). Known host plants
are grasses in the genera Andropogon. Calamagrostis. Schizachvrium and Sorghastmm
(Hamilton, pers. comm.; Panzer et al.., 1995). The widespread Chlorotettix unicolor is
transcontinental (Beime, 1956; Delong, 1948) and is associated with a variety o f habitats, from
w et to dry, pristine native to highly degraded (Bess, pers. obs.; Delong, 1948; Panzer, pers.
56
comm.). In 1991 and 1992,1 swept numerous specimens o f C. Unicolor from isolated patches o f
Agropyron smithii and A. spicatum on the STCO/BOGR grasslands surrounding Logan,
Montana. Both nymphs and adults were collected from these grasses, further confirming their use
as hosts by this leafhopper. From an analysis o f the 1988 and 1991 data, C. unicolor was found to
be most common in the FEID/AGSP grasslands (Table 5), where its abundance correlated with
percent cover by native Agropyron spp. (Table 11). This leafhopper was also swept in low
numbers from the replanted grasslands, where it may be using introduced species o f Agropyron as
hosts.
Diplocolenus configuratus
Diplocolenus configuratus is native to Montana and was very abundant in the Brin/Mesa
habitats (Table 5). This leafhopper also occurred in the FEID/AGSP patches and I collected
several specimens o f D. configuratus above treeline on Mount Blackmore and along the crest of
the Bridger Mountains in 1991 and 1992. Delong (1948) reported this species as occurring in
association with Canada bluegrass fPoa compressa), a non-native, European grass species found
throughout the East and Midwest (Hitchcock, 1951). I have collected D. configuratus from Poa
compressa and P. pratensis in a variety o f habitats across the Midwest, including my lawn in
northwest Indiana. The abundance o f D. configuratus at the twelve patches correlated highly with
percent cover by Poa spp. (Table 11).
57
Doratura stvlata
D oratura stvlata is a European leafhopper that has spread westward from its original
introduction in the Great Lakes region (Hamilton, 1972). This species is extremely abundant in
Poa-dominated habitats such as pastures and lawns in the Midwest (Bess pers. obs.; Delong,
1948). Beime (1956) stated that D. stvlata probably feeds on grass roots in marshes and
meadows. Abundance o f this species at the twelve patches correlated strongly with percent cover
by Poa spp. (Table 11). This leafhopper was most abundant in the Brin/Mesa association,
although it occurred in lesser numbers in the FEID/AGSP association (Table 5).
Dorvcephalus platvrhvnchus
Dorvcephalus platvrhvnchus is reported to feed on wild rye IElymus spp.) and is known to
use native and introduced species o f Agropvron (Bess, pers. obs.; Hamilton, pers. comm.; Panzer
et a l, 1995). This leafhopper’s use o f native Agropyron species was confirmed by significant
correlations between its abundance and percent cover by Agropvron smithii and A. spicatum
(Table 11). Gould (1947) considers all o f the native North American grass species previously
assigned to Agropyron to be included in Elymus. meaning that D. platvrhvnchus is probably
monophagous on Elymus.
Endria inimica
Endria inimica is a native, transcontinental leafhopper species occurring in a variety o f
habitats (Beime, 1956; Delong, 1948). It appears to prefer Poa pratensis as a host (see Beirne,
1956) and I have collected large numbers of both E. inimica and D. stvlata from Poa pratensisdominated fields in the Midwest. The abundance o f this leafhopper at the twelve patches
correlated strongly with percent cover by Poa spp. (Table 11).
58
Flexam ia ab b rev iatta and F. flexulosa
Flexamia abbreviata and F. flexulosa are known to feed exclusively on the grass genus
Bouteloua. preferring B. gracilis and B. hirsutus (Whitcomb and Hicks, 1988). Both Flexamia
species were recorded only from the STCO/BOGR plant assemblage (Table 5). Abundance o f
Flexamia flexulosa correlated strongly with percent cover o f its only known host, Bouteloua
gracilis. The abundance ofF . flexulosa also, correlated significantly with percent cover o f Stipa
comata, which occurs in the same habitat, but this grass is not considered a host. Hamilton (pers.
comm.) stated that few leafhopper species appear to specialize on Stipa comata as a food plant.
Both species o f Flexamia reach the northwestern limit o f their respective ranges in western
Montana and Alberta (Whitcomb and Hicks, 1988).
H ecalus spp.
Hecalus leafhoppers occurred most abundantly in the Agcr/Mesa grasslands, especially at
17b (Table 4). Males were identified as belonging to both Hecalus major and H. viridis. but
Whitcomb and associates (1996) consider the taxonomy o f this group to be in question, thus
rendering specific identifications questionable. Both Hecalus major and H. viridis are considered
moderately remnant dependent in Illinois (Panzer et a!, 1995), where H. major is associated with
wet prairie containing its host, Calamagrostis canadensis (Delong, 1948; Panzer et al., 1995).
Hecalus viridis is considered a species o f xeric prairie in Illinois, where it is reported to feed on
“native grasses” (Panzer et a l, 1995).
Hecalus leafhoppers are certainly not remnant dependent in Gallatin County, Montana,
where they were found to be fairly common in native and replanted patches. These leafhoppers
were most abundant in the Agcr/Mesa assemblage. Abundance of Hecalus spp. showed positive
59
correlations with cover by Agropvron grasses (Table 11), their suspected host plants in Gallatin
County. Adults o f these leafhoppers appeared in large numbers in the late spring (late May/early
June) samples, while the mid-summer (late July) samples were dominated by nearly full grown
nymphs. This suggests that Hecalus leafhoppers produce a brood prior to when A. cristatum
becomes semi-dormant in the heat o f August, allowing them to successfully use this grass as a
host.
L atalus missellus
Latalus missellus is reported to occur throughout the northern United States, Canada and
Alaska (Beirne, 1956; Delong, 1948). In my study, this leafhopper was found almost exclusively
in the Brin/Mesa association (one specimen collected at patch 26 A in the FEED/AGSP
association), where its abundance correlated strongly with percent cover by Poa spp. (Table 11).
I have also collected L. missellus from Poa compressa and P. pratense in Indiana.
Q rocastus labeculus and O. perpusilus
The two species o f Orocastus are reported to feed on Stipa comata (Hamilton, pers.
comm.) and their abundance correlated strongly with percent cover by this species (Table 11).
Both leafhoppers were most abundant at patch IOA (Table 6), which had the greatest percent
cover by Stipa comata (Table 2). Orocastus perpusillus was most abundant in the spring samples
(May and June), when this cool season grass is in full bloom and at a maximum growth state.
Orocastus labeculus peaked in abundance roughly one month later when this grass species was
setting seed. It has been hypothesized that leafhopper species abundance is closely tied to
seasonal growth flushes in their host plants, especially during flowering and fruit set (Waloff,
1980; Whitcomb et al.„ 1987).
60
Conclusions Concerning LeafhopperZHost Plant Associations
The abundance o f several leafhopper species collected from grassland patches in the
Gallatin Valley correlated closely with percent cover o f known host plants, thus supporting the
hypothesis (for some species) that the distribution o f leafhoppers is correlated with the
distribution o f their host plants. Further observational and experimental studies to determine
exact leafhopper-host plant associations would help interpret the distribution o f additional
leafhopper species among the four plant assemblages. Such studies would also help determine
whether some o f the weaker leafhopper/host plant correlations observed in this study were
actually a result o f host plant specificity or were a result o f some other limiting factor(s) (e.g. if
the leafhopper had similar abiotic requirements as the plant but did not actually feed on it).
Leafhopper Taxa occurring Primarily in the Native Plant Assemblages
Several leafhopper taxa were recorded exclusively (or primarily) in the native grassland
patches. These include Chlorotettix unicolor. Deltocephalus valens. Mocuellus caprillus.
Orocastus perpusillus, Rosenus cruciatus. Stenometopielus cookei. and Xerophloea viridis.
Chlorotettix unicolor and Xerophloea viridis were most abundant in the FEID/AGSP assemblage
while the remaining species were most abundant in the STCO/BOGR patches. Chlorotettix
unicolor and Mocuellus caprillus are native Agropyron feeders in Gallatin County (Bess, pers.
obs ), although C. unicolor is associated with Andropogon and Calamagrostis grasses in the
Midwest (Bess, pers. obs.). In 1991 and 1992,1 collected numerous individuals o f M- caprillus
from Agropvron smithii and A. spicatum on the grasslands surrounding Logan, Montana, in
association with the leafhoppers Auridius helvus and Chlorotettix unicolor.
61
Orocastus
perpusillus is a Stipa comata specialist throughout southern Canada and the northwestern United
States (Hamilton, pers. comm.). Xerophloea viridis is reported to feed on Aristida grasses in the
Midwest (Delong, 1948). The food plants o f the remaining species are unknown for Gallatin
County.
Deltocephalus valens and Rosenus cruciatus were both most abundant in the 1988 samples
from the STCO/BOGR patches, but were almost entirely absent from the 1991 samples. The
food plants o f these leafhoppers are unreported, but assumed to be grasses. I have swept R.
cruciatus from Koeleria cristata in oak savanna’s o f northwest Indiana, where this leafhopper is
considered to be very rare (Panzer et al., 1995). Delong (1948) considered R. cruciatus to be a
prairie species. N ot much is known about Deltocephalus valens. but other Deltocephalus species
in the Midwest are considered host specific and/or remnant dependent (Bess unpub. data; Delong,
1948; Panzer et al., 1995).
Stenometopiellus cookei appears to overwinter as an adult and is among the first
leafhoppers to appear in the spring, usually in association with the leafhopper Hardya dentata and
nymphs o f various Athvsanella spp. (Bess, pers. obs.). I collected many individuals o f both S.
cookei and H. dentata in M arch and April, from grasslands dominated by Agropvron smithii. A.
spicatum, Stipa comata; and S. viridula near the towns o f Logan and Three Forks. These two
leafhoppers belong to genera that are primarily Eurasian in distribution, with disjunct species (S.
cookei. H. dentata and the closely related H. voungif in the western United States and Canada
(Hamilton, 1983).
62
L eafh o pper T axa occurring P rim arily in the R eplanted P lan t Assemblages
An interesting phenomenon observed during this study was the occurrence o f several
leafhopper species almost exclusively in the replanted patches. This grouping includes
AmblysellUs grex. Athvsanus argentarius. Balclutha punctata. Diplocolenus configuratus.
D oratura stylata. Elvmana circius. Endria inimica. the genus Hecalus. Latalus missellus, and a
species o f Lonatura. All o f these species, with the exception o f Lonatura. were recorded almost
exclusively from the Brin/Mesa patches. The Lonatura females were recorded from patch 16b in
the Agcr/Mesa assemblage.
Athvsanus argentarius and Doratura stvlata are European leafhopper species introduced
into the Midwest and having since spread westward to at least Montana. Both are considered
general grass feeders, although their abundance at the twelve patches correlated significantly with
percent cover by Poa spp. The leafhoppers Diplocolenus configuratus. Endria inimica. and
Latalus missellus are native species associated with northern meadow and pasture communities in
southern Canada and the northern United States. All three are considered Poa feeders (Beime,
1956; Bess, pers. obs.; Delong, 1948) and their abundance across the twelve patches correlated
significantly with percent cover by these grasses.
Balclutha puncatata is another widespread leafhopper species occurring in a variety o f
grasslands, pastures and agricultural situations throughout temperate regions o f the world
(Beirne, 1956; Delong, 1948; Hamilton, 1983; WalofF,, 1973). Whitcomb et al. (1994) consider
this leafhopper to be a general grass feeder in North America, preferring species in the tribe
Chloridoideae. This is an interesting observation, as no chloridoid grasses were found at patch
2 1A, where this leafhopper was collected in very large numbers (Table 6). Poa pratensis and
63
Bromus inermis were the only grass species found in the sampling plots at 21b, occurring very
frequently and providing relatively extensive cover (Table 2). Both grasses belong to the tribe
Poeae. Therefore, it appears that B. punctata may be utilizing these non-chloridoid grass species
as hosts in southwest Montana.
Elvmana circius is a species new to the leafhopper fauna o f the United States, being
previously known from populations in the mountains o f Alberta and British Columbia (Hamilton
and Chiykowski, 1985). It is most likely that E. circius is feeding on Agropvron spp. (=Elymus,
following Gould, 1947) in southwest Montana, as members o f the genus Elvmana are reported to
feed predominantly on grasses in the genus Elymus (Hamilton and Chiykowski, 1985; Delong,
1948). Hamilton and Chiykowski (1985) also report E, circius from Bromus and “other grasses” .
In the Brin/Mesa association, this leafhopper was recorded only from patches 25B and 26B. Both
patches have (in addition to Bromus inermis) a high frequency and percent cover o f Agropvron
intermedium, a potential host plant. Several E. circius were also recorded at patch 26A in the
FEID/AGSP association, where they may be feeding on Agropvron spicatum.
Beime (1956) states that Lonatura leafhoppers are associated with grasslands and the host
plants are probably grasses. The specimens collected from 16b are pale yellowish and unmarked,
putting them close to L. crocea (Beirne, 1956). Delong (1948) associated L. catalina with the
grass Aristida gracilis in Illinois. Stipa is closely related to Aristida and represents a potential host
for the Lonatura species recorded at 16b, where S. comata was fairly frequent (Table 2).
64
Leafhopper Taxa with Wide Distributions in the Gallatin Valley (but no well-defined plant
assemblage or host plant preferences)
Several o f the more abundant leafhopper taxa displayed no obvious association with any
one plant assemblage. These include members o f the genus Auridius. Colladonus geminatus.
Cuema striata. Hebecephalus rostratus. Macrosteles quadrilineatus. and Psammotettix lividellus.
The Auridius leafhoppers appear to be using species o f Agropvron as hosts (Bess, pers. obs.;
Hamilton, pers. comm ), although they were conspicuously rare in the samples from the
Agcr/Mesa patches. The BrinZMesa collections were nearly all (78 o f 85) from patch 26b which
had abundant cover o f Agropvron intermedium, a replanted species native to Montana. It is
possible that species o f Auridius prefer native Agropvron as hosts.
Colladonus geminatus and Cuema striata are widely distributed throughout the United
States and southern Canada, where they are reported to feed on a variety o f herbaceous plants,
trees and shrubs (Beime, 1956; Delong, 1948). Colladonus geminatus was most abundant in the
patches at the more mesic end o f the valley, especially in the BrinZMesa assemblage. This
leafhopper is reported to feed on a variety o f trees and shrubs (Beime, 1956). Cuerna striata was
most abundant at the more xeric end o f the valley, being somewhat more common in the
STCO/BOGR assemblage. The genus Cuema is primarily boreal and montane in distribution
(Beime, 1956), with Cuerna striata occurring throughout Canada and the United States (Beime,
1956; Delong, 1948). Delong (1948) reports C. lateralis (= striata Walker) to be “abundant in
pastures and herbaceous growth in cutover areas” in Illinois. Its food plants in Montana are
unknown.
65
Psammotettix lividellus is a circumpolar species occurring across Canada and the northern
United States (Beirne, 1956; Hamilton, 1983). In this study, P. lividellus occurred across all
patches, but was most abundant in the replanted grasslands. In 1991 and 1992,1 collected this
species in a variety o f native and non-native plant assemblages throughout Gallatin County.
Given its broad distribution, it is probable that P. lividellus is a general feeder on a variety of
grasses. Hebecephalus rostratus is a grassland species occurring in southwestern Canada and
northwestern United States (Beirne, 1956). It was found in nearly all o f the patches sampled, but
was most abundant in the native plant assemblages (esp. STCO/BOGR). Its food plants are
unknown, but assumed to be grasses.
Notes on the Rarer Leafhopper Species Recorded in this Study
Nearly half (20) o f the leafhopper species recorded in this study were represented by fewer
than ten individuals. Some o f these may have hosts that are extremely patchy in distribution,
making the leafhoppers difficult to sample in a systematic fashion. Still others may be highly
transient, blowing in on air currents, only to leave for new grasslands with the next weather
system. A final subset may have behaviors that make them unlikely to be collected with a sweep
net. The following is a discussion of some of the more unusual leafhopper species collected
during this study and possible explanations for their rarity in the samples.
Frigartus frigidus is reported to feed on Artemisia frigida (Beirne, 1956). In 1991 and
1992,1 swept several individuals from grasslands containing this plant near patch 9A. The area
consisted o f protected canyons with vegetation dominated by Agropvron smithii and Stipa
viridula. This leafhopper is large bodied and probably not especially mobile. Therefore, it may be
66
difficult to collect unless its foodplants occur in the immediate vicinity o f the sampling area and
are known and recognized by the sampler.
Leafhoppers o f the genus Prairiana appear to be partially Subterranean and at least
somewhat nocturnal (Bess, pers. obs.; Hamilton, 1994 pers. comm.). Their host plants are
Unknown, although Hamilton (1994) suspects sages f Artemesia spp.) for some o f the western
species. I have taken the eastern Prairiana kansana and P. angustens in habitats dominated by
species o f Andropogon grasses and various Asteraceae, but with no Artemisia spp. present (Bess,
unpub. data). Other leafhoppers in the Gyponinae (the subfamily to which Prairiana belongs) are
primarily tree and shrub feeders (Delong, 1948; Hamilton, 1996 pers. comm.). Prairiana
leafhoppers are usually rare in sweep net samples, although I observed more than three dozen
individuals o f P. angustens at an ultraviolet light on one night in-Kentucky. This species
overwinters as an adult in Kentucky where it is known from one grassland preserve (Bess unpub.
data).
Leafhoppers in the genus Commellus are known to feed on Agropvron and Stipa grasses
in the Midwest (Panzer et al., 1995). In my study, C. sexvitattus had a peculiar distribution, being
recorded in essentially equal numbers from both a native Stipa-dominated (IOA) and a non-native
Agropyron-dominated patch (25b). It is possible that C. sexvitattus feeds on grasses from both
genera in southwest Montana.
Neocolidia tumidifrons belongs to a genus distributed primarily in the American west
(Delong, 1948). It is a widely distributed species, occurring from the Pacific east through the
Great Lakes States (Delong, 1948). In the Midwest, it is associated with mesic habitats with rich
67
herbaceous growth (Bess, pers. obs.; Delong, 1948). It is likely more abundant in mesic plant
assemblages, possibly ones that were not sampled in this study.
Mesamia coloradensis is a species characteristic o f prairies throughout the Midwest and.
northern Great Plains states (Delong, 1948). Members o f the genus Mesamia are thought to feed
exclusively on plants in the family Asteraceae and M. coloradensis has been reported as feeding
on Artemisia frigida (Beime, 1956). Related species o f Mesamia feed on the genus Helianthus in
the Midwest (Bess, pers. obs.; Delong, 1948; Panzer et al., 1995).
Hardya dentata was recorded only from samples taken in the Agcr/Mesa grasslands
(Tables 3 and 4). Beirne (1956) reported this species from southern Saskatchewan and stated that
it may live in litter or among plant roots. I collected many individuals o f H dentata and
Stenometopielus cookei in March and April, from grasslands dominated by Agropvron smithii. A.
spicatum. and Stipa viridula near the towns o f Logan and Three Forks. These two leafhoppers
belong to genera that are primarily Eurasian in distribution, with disjunct species (S. cookei. H.
dentata and the closely related H. voungi) in the western United States and Canada (Hamilton,
1983).
Idiodonus aurantiacus was represented by two females taken in June o f 1988 from site Tb
in the Brin/Mesa assemblage (Appendix B). Beirne (1956) reports I. aurantiacus as being
widespread, but local, in western Canada. Host plants for this genus are unreported, although
species o f the closely related genus Colladonus feed on a variety of herbaceous .plants, trees, and
shrubs.
68
Notes on Some of the Pooled Leafhopper Genera
Several leafhopper taxa were identified only to genus because o f taxonomic difficulties
discussed in the Materials and Methods section. The genus Aceratagallia was discussed
previously in this section, but three additional genera need to be addressed. These are
Athvsanella. Dikraneura. and Sorhoanus. Based on the male specimens examined in 1988, seven
Athvsanella species (acuticauda. attenuata, occidentalis. robustus. sinuata. terebrans, and utahna)
were recorded from the STCO/BOGR plant assemblage (Appendix A). Athvsanella sinuata is
reported to be a Bouteloua gracilis specialist, while A. occidentalis feeds primarily on this grass
and other warm-season species (Whitcomb, et al., 1994). Athvsanella attenuata is reported to
feed on wheatgrass (Blocker and Johnson, 1990) and A. acuticauda is reported to feed on
bluegrass (Poa spp:). The remaining Athvsanella species are reported to be more general on other
warm-season grasses (Blocker and Johnson, 1990; Whitcomb et al., 1994).
Delong (1948) reports ~60 species o f Dikraneura as occurring in the Nearctic region.
Beime (1956) reports nine o f these from Canada. Male specimens o f this genus were identified as
D. shoshone. but many samples contained mostly females. Dikraneura leafhoppers are small,
widespread species occurring in a variety o f grassland types throughout Gallatin County (Bess,
pers. obs.). They were most abundant in the Brin/Mesa association.
The genus Sorhoanus contains five described species in northwestern N orth America
(Beime, 1956), three o f which (debilis. flavovirens and orientalist were identified in samples taken
during this study (Appendix A). These leafhoppers were most abundant in the FEID/AGSP
patches, where they were the most numerous leafhopper collected (Table 5). In contrast to their
abundance in the FEID/AGSP plant assemblage, samples from the STCO/BOGR patches
69
fluctuated widely in abundance o f these leafhoppers, with 16A recording the lowest number o f
individuals (Table 6). This STCO/BOGR patch is located south o f Three Forks, some distance
from any o f the local mountain ranges which may contain the preferred habitats for these
leafhoppers. Beime (1956) reports Sorhoanus leafhoppers from northern grassland, arctic, and
mountainous habitats, reaching their southern distribution limit in the northern United States.
Whitcomb and associates (1987a) hypothesize that these leafhoppers are associated with pooid
grasses. My data suggest that, o f the four grassland types surveyed, these leafhoppers prefer
those at higher elevations (Table 5).
70
SAMPLING EFFORT AND ADDITIONS TO THE
LEAFHOPPER FAUNA OF MONTANA
Sampling Effort
A major difficulty in this study was the large number o f specimens encountered and the
physical similarity o f many o f the species examined. Initially, it was my intention to compare
leafhopper samples from each o f the 39 grassland patches studied by Kemp et al. (1991). This
goal proved unattainable following initial counting and identification o f samples from the twelve
patches used in the analysis. At a rate o f 20,000 specimens/12 sample patches/year (average), 39
sample patches would produce at least 65,000 leafhoppers/year, for a minimum o f 260,000
specimens for a four-year study.
Therefore, I was forced to reduce the number o f patches and years to a manageable level.
This reduction in the number o f patches decreased the strength o f conclusions that could be
drawn from the analyses. A greater number o f patches and a greater number o f samples per patch
would have allowed for the calculation o f mean and variance abundance statistics for each. This
would have allowed for a more detailed comparison o f within patch, and within plant assemblage,
variance for individual species.
Additional limitations, when working with leafhoppers, involve the method used to collect
specimens. Over the years, a variety o f techniques have evolved, falling into three primary
categories; sweep netting, suction sampling and “passive collectors” (i.e. sticky traps, pan traps,
flight intercept traps). Sweep netting and suction sampling are the most commonly used methods
for sampling leafhopper assemblages in grassland ecosystems (Blocker and Reed, 1976; Brown et
71
al., 1992; Cherrill and Rushton, 1993; Cwikla and Blocker, 1981; Morris, 1973, 1981a-b, 1983;
Novotny, 1995; Waloff, 1973, 1980; WalofFand Solomon, 1972; Whitcomb et al., 1994).
Blocker and Reed (1976) found that, in a tallgrass prairie remnant, similar numbers o f leafhopper
species were collected using the two methods, but abundance varied greatly, with the D-vac
suction samplers collecting the greatest number o f individuals.
Therefore, in studies attempting to determine subtle similarity (or differences) between
habitat patches, suction samplers may be more appropriate. In tallgrass prairie and wetland
communities, where vegetation often exceeds shoulder height, sweep netting can be very difficult
and the use o f suction samplers may be more advantageous. However, this method is very time
consuming and labor intensive, and may be inappropriate for studies in remote regions where
equipment would need to be carried long distances. Therefore, the methods used for sampling
leafhoppers depend on the characteristics o f the vegetation at hand.
Given that the grasslands covered in this study never exceeded I m in height, it can
generally be assumed that sweep netting was an appropriate method for sampling leafhoppers. In
an eight-year study o f the grasslands o f Kentucky, with vegetation o f similar stature, greater than
95% o f the local leafhopper fauna was collected by sweep net in the first two years (Bess, unpub.
data). Panzer (pers. comm.) has reported similar results in Illinois. Finally, sticky traps and pan
traps can also be highly effective means o f sampling Cicadellidae, but specimens can be difficult to
process and identify.
As a final discussion o f sampling effort and its bearing on this study, Novotny (1994), in a
four-year study o f 62 sites in central Europe, examined 255 populations o f 101 leafhopper species
represented by -105,000 individuals. In the present two year study o f 12 patches in southwestern
72
Montana, over 44, 000 individuals o f at least 64 leafhopper species were observed, nearly twothirds o f Novotny’s number o f species on one-twentieth o f the sample sites. The large number o f
individuals per sample and the relatively high diversity o f species suggests that the sweep net
sampling method used in this study was appropriate.
The lists o f leafhoppers reported here probably do not represent a complete catalog o f all
leafhopper species potentially present at the sites sampled. In fact, 15 o f the 56 taxa collected in
1988 did not appear in the 1991 samples. However, only one o f the 15 made up more than one
percent o f the individuals collected within a plant assemblage (i.e. Rosenus cruciatus constituted
3.6% o f the leafhoppers collected in the STCO/BOGR patches). Only five o f 54 taxa collected in
1991 had not appeared in the 1988 samples, although only one o f those five made up more than
one percent o f the individuals collected within a plant assemblage (i.e. Auridius auratus comprised
1.7% o f the leafhoppers from the FEID/AGSP patches). The differences observed in leafhopper
abundance between the sampling years may have been a result o f I) the effect o f sampling method
on the sampler’s ability to collect rare or patchily distributed species, 2) extirpation o f some o f the
15 leafhopper species that had been present in 1988, 3) migration o f new leafhopper species into
patches after 1988, or 4) differences in yearly weather patterns. The nature o f my data make it
impossible to test these hypotheses.
The differences observed between samples from the two years suggest that more intensive,
perhaps longer-term, sampling would be required to obtain a complete catalog o f the species
present within the plant assemblages covered in this study. However, for several reasons, the
differences observed between the two sampling years probably do not weaken the major
conclusions o f this study. First, the correlation analyses for each year were not affected by the
73
absence o f certain species, because only the leafhoppers collected in each year were used for that
year’s analysis. Ifth e absent species had also been used, the correlations would have been
artificially inflated because o f multiple zero-zero data pairs. Second, few o f the species observed
only in one year were major components o f the leafhopper fauna at their particular plant
assemblage. Third, the species found only in one year were irrelevant to the analysis of
correlations between leafhopper species abundance and percent cover o f potential host plants.
Additions to the Leafhopper Fauna of Montana
Fox (1924) recorded 52 species o f leafhoppers in 27 genera from Montana, primarily from
areas around Bozeman. In contrast, at least 67 distinct leafhopper taxa in 5 1 genera were
identified from the 1988 and 1991 samples used in my study (Table 12 and Appendix B). By
comparison with Fox’s species list, 54 o f these leafhopper taxa are new to the state o f Montana.
Several o f these (such as Athvsanella acuticauda. A. occidentalis. A. sinuata. Flexamia abbreviatta
and F. flexulosal have been reported from Montana in the literature (Blocker and Johnson, 1980;
Whitcomb et al., 1994). This brings the total state leafhopper fauna to 112, a number that could
easily be increased through further survey effort.
74
Table 12. The Leafhopper Fauna of Montana 1
L E A F H O P P E R S P E C IE S
Aceratagallia cinerea
Aceratagalha novella
Aceratagallia sanguinolenta
Aceratagallia uhleri
Agallia quadripunctata
A m blyseU us grex
A th ysa n ella acuticauda
A th ysa n ella attenuata
A th ysa n ella occidentalis
A th ysa n ella robusta
A th ysa n ella sin u a ta
A th ysa n ella terebrans
A th ysa n ella utahn a
A th y sa n u s argentarius
Auridius auratus
A u rid iu s helvus
A u rid iu s ordinatus
Balclutha abdominalis
Balclutha confusa
Balctutha impicta
Balclutha punctata
Bythoscopus rufoscutellatus
Chlorotettix unicolor
C icadula quinquinotata
Colladonus bolli
Colladonus fasciaticollis
Colladonus geminatus
Colladonus montanus
Commellus comma
C om m ellus sem icolon
C om m ellus sexvita ttu s
C uerna striata
D eltocephalus valens
B ess
*
*
Diplocolenus conjiguratus
Doratura stylata
Dorycephalus platyrhynchus
Draeculacephala manitobiana
Draeculacephala mollipes
*
*
*
*
*
sp.
E n d ria rotunda
*
*
*
*
*
$
*
*
*
*
*
*
*
$
*
*
*
*
*
*
*
*
*
*
*
*
*
♦
Id iodonus aurantiacus
L aevicephalus sp.
*
Latalus misellus
Limotettix spp.
*
*
*
*
L o n a tu ra sp.
*
*
*
Erythroneura comes
*
*
M acrosteles quadriUneatus
M esa m ia coloradensis
M o cu ellu s caprillus
*
*
*
*
Neokolla hieroglyphica
Neokolla gothica
*
*
*
N eocolidia tu m id ijro n s
N esosteles neglectus
NorveU na se m in u d a
*
*
O rocastus labeculus
O rocastus p erp u sillu s
P araphlepsius occidentalis
P in u m iu s areatus
P rairiana cinerea
P rairiana subta
P sa m m o tettix Uvidellus
*
*
*
*
*
*
*
Psammotettix striatus
*
R o sen u s cruciatus
*
Scaphytopius acutus
*
*
Sorhoanus debilis
So rh o a n u s Jlavovirens
So rh o a n u s orientalis
Sten o m eto p ielu s cookei
S treptanus co n jin is
T exananus cu m u la tu s
*
T o ta l:
75
*
*
Xerophloea major
Xerophloea viridis
Xestocephalus pulicarius
1 E n trie s in B O L D a re n e w s p e c ie s re c o rd s f o r M o n ta n a
*
*
*
*
Macropsis sordida
Macropsis viridis
Scaphytopius Jrontalis
*
*
*
*
*
*
*
*
Norvelina tenella
Oncometopia lateralis
Uncopsis distinctus
*
*
*
*
*
Hecalus viridis
Helochara communis
Idiocerus lachrymalis
Idiocerus snowi
Idiocerus suturalis
*
*
*
*
HecaUis m a jo r
*
*
*
Fox
*
Hebecephalus signatij'rons
E lym a n a circius
E m poasca
B ess
*
*
*
*
E lexa m ia abbreviata
E lexa m ia Jlexu lo sa
E orcipata Ioca
E rigartus frig id u s
H ardya dentata
H ebecephalus rostratus
Empoasca alboscripta
Empoasca aspersa
Empoasca Jlavescens
Empoasca mali
Endria inimica
L E A F H O P P E R S P E C IE S
Exitianus exitiosus
Gypona bimaculata
Dikraneura cameola
Dikraneura Jieberi
Dikraneura mali
D ikratieura sh o sh o n e
D ikrella s p .
Fox
*
*
*
67
52
SUMMARY
1. Leafhopper samples from the three patches within each o f the four plant assemblages
were found to correlate significantly with one another using the combined data. The
STCO/BOGR patches were found to be the most closely associated o f the four types sampled.
However, there were also a number o f significant correlations between patches from different
plant assemblages. These between-plant assemblage correlations usually occurred when the
patches were close spatially and contained similar vegetative components (e.g. the Agropvrondominated habitats).
2. In addition, leafhopper samples from patches within some o f the plant assemblages
(e.g. the FEID/AGSP and Brin/Mesa habitats) did not correlate closely with one another in either
one, or both, o f the sampling years. The BrinZMesa patches did not correlate significantly with
any o f the other patches in analysis o f the combined data.
3. The abundance of selected leafhoppers was strongly correlated with percent cover by
known or suspected food plants. M ore intensive studies are needed to determine host plant
associations for most of the leafhopper species encountered in this study. These associations are
crucial to formulating an understanding o f the habitat requirements o f these insects.
4. Cluster analyses, based on Goodman-Kruskal’s gamma coefficients, produced many
spurious associations when comparing patches using an individual year’s data. Cluster analysis o f
the combined year’s data, however, produced associations supported by the correlation analysis
and patterns observed in the raw data. Samples from the low elevation native grasslands
(STCO/BOGR) were found to be the most closely associated and were also closely associated
76
with those from the mid-elevation native patches (FEID/AGSP). Given the spurious associations
between patches using the single years’ data, Goodman-Kruskal’s Gamma analysis should be used
with great caution in situations where the data set conntains a wide range o f values and numerous
missing data points.
5. Leafhopper samples from patches 26A, 25b and 26b formed a distinctive outlying
group unrelated to those from the remaining nine sites. The three sites contained many leafhopper
species that were never, or only rarely, recorded from the other patches.
6. The results indicate that grasslands replanted to non-native plant species often harbor
large numbers o f a few, widespread leafhopper species. Two o f those observed in this study,
Athysanus argentarius and Doratura stylata. are not native to North America and have spread
rapidly from their sites o f introduction in eastern Canada. Doratura stvlata was, by far, the most
abundant leafhopper collected in this study and 99 percent o f the individuals came from the
replanted Bromus inermis/Medicago sativa grasslands. Many o f the other wide-ranging
leafhopper species observed in these man-made habitats are also considered agricultural pests and
likely impart a tremendous stress on the plants they feed on. Some are also known vectors o f
plant diseases.
7. A total o f 54 new leafhopper species were added to the known fauna o f Montana and
this number could likely be increased through further study.
77
B IB LIO G R A PH Y
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Vegetation structure and microhabitat selection in grasshoppers (Orthoptera:
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Census data o f populations o f some leafhoppers
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83
A PPEN D IX A: 1988 AND 1991 LE A FH O PPE R PH EN O LO G Y DATA
F O R TH E TW ELV E PA TCH ES
84
A ppendix A.
1988 Leafhopper Phenology Data.
The Stipa comata/Bouteloua gracilis Plant Assemblage
LEA FH OPPER
S P E C IE S
Patch 7A
Patch IOA
Patch 16A
Sampling Dates
Sampling Dates
Sampling Dates
Orocastus perpusillus
432
6
Mocuellus caprillus
145
4
11
43
3
7
Rosenus cruciatus
91
3
4
14
I
3
Auridius helvus
63
Sorhoanus debilis
53
Deltocephalus valens
31
Hebecephalus rostratus
11
Athysanella utahna
8
Colladonus montanus
8
Athysanella occidentalis
7
I
2
7
25
Aceratagallia spp.
5
86
Dorycephalus platyrhynchus
4
2
Endria rotunda
3
2
Athysanella acuticauda
I
I
Colladonus geminatus
216
156
234
2
46
12
38
135
6
157
57
20
121
68
18
108
22
74
8
12
16
Psammotettix lividellus
6
8
20
2
I
60
10
6
2
2
40
72
121
2
7
Hecalus spp.
2346
9
Dikraneura spp.
Prairiana cinerea
120
1788
Dorycephalus platy.
Macrosteles quad.
TOTAL
6/3/88 7/19/88 8/23/88 6/3/88 7/19/88 8/23/88 5/26/88 7/18/88 8/22/88
149
27
20
13
16
I
370
23
I
3
3
I
2
3
2
2
3
10
10
16
I
2
2
5
6
I
7
Orocastus labeculus
44
5
45
84
Cuem a striata
42
6
2
8
Flexamia abbreviata
3
Flexamia JlexuIosa
I
Frigartus frigidus
I
Xerophloea viridis
21
199
58
3
29
2
I
32
3
I
I
I
Stenometopiellus cookei
48
48
Commellus sexvittatus
15
15
Athysanella terebrans
13
13
Pinumius aeratus
2
2
Athysanella sp.
I
I
Empoasca sp.
I
I
Streptanus confmis
I
I
2
Prairiana subta
Athysanella robusta
TOTAL (N)
NUMBER O F SPECIES
2
I
I
883
218
176
2170
62
133
533
170
2
4347
21
15
6
24
8
9
14
7
2
29
85
A ppendix A.
1988 Leafhopper Phenology Data.
T he Festuca idahoensis/Agropyron spicatum Plant Assemblage
LEA TH O PPER
SPECIES
Sorhoanus debilis
Rosenus cruciatus
Psammotettix lividellus
Dikraneura spp.
Macrosteles quadrilineatus
Deltocephalus valens
Orocastus perpusillus
Athysanella acuticauda
Hebecephalus rostratus
Aceratagallia spp.
Hecalus spp.
Dorycephalus platy.
Orocastus labeculus
Xerophloea viridis
Athysanella sinuata
Diplocolenus con/iguratus
Chlorotettix unicolor
Patch 21A
Patch 2 SA
Patch 26A
Sampling Dates
Sampling Dates
Sampling Dates
TOTAL
6/2/88 7/18/88 8/22/88 6/2/88 7/18/88 8/22/88 6/16/88 7/18/88 8/22/88
223
44
34
14
14
11
10
9
9
7
7
8
159
18
79
15
13
3
I
I
9
I
5
7
2
34
15
I
7
5
7
38
2
3
2
2
I
5
2
8
9
123
63
7
2
2
49
28
22
3
I
2
10
I
65
10
9
4
4
4
36
27
34
20
no
2
271
25
6
I
5
I
25
2
16
I
I
66
114
39
20
Macrosteles quad.
Commellus sexvittatus
Collaadonus geminatus
615
3
I
Endria inimica
Athysanella utahna
225
I
9
I
7
21
4
187
69
20
20
23
15
I
13
3
Mesamia coloradensis
I
I
Mocuellus caprillus
I
58
4
14
I
9
4
Dorycephalus platyrhynchus
7
Neocolidia tumidifrons
Athysanella occidentalis
5
20
2
9
5
7
2
Cuema striata
Auridius helvus
Elymana sp.
6
2
9
I
3
31
I
31
I
I
Limotettix dasidus
Limotettix kryptus
I
10
10
2
Laevicephalus sp.
2
2
Norvelina seminuda
2
2
Athysanella attenuata
I
I
Frigartus frigidus
Texananus spp.
I
I
I
Ballana veruta
2
I
TO TA L (N)
391
263
59
342
129
8
405
191
6
1794
NUMBER O F SPECIES
16
14
9
17
12
4
17
15
3
33
86
A ppendix A.
1988 Leafhopper Phenology Data.
T h tA g ro p yro n cristatum/Medicago sativa Plant Assemblage
LEA FH OPPER
S P E C IE S
Dikraneura spp.
Psammotettix Hvidellus
Aceratagallia spp.
Rosenus cruciatus
Colladonus geminatus
Hecalus spp.
Sorhoanus debilis
Orocastus perpusillus
Deltocephalus valens
Auridius helvus
Hebecephalus rostratus
Dorycephalus platy.
Orocastus labeculus
Diplocolenus configuratus
Elymana sp.
Lonatura sp.
Prairiana cinerea
Patch 7b
Patch 16b
Patch 17b
Sampling Dates
Sampling Dates
Sampling Dates
6/3/88
89
41
15
15
10
9
8
7
6
TOTAL
7/21/88 8/23/88 5/26/88 7/21/88 8/24/88 5/26/88 7/20/88 8/25/88
2
1024
374
I
35
113
4
3
5
14
31
I
9
5
Il
360
6
2
137
515
21
1458
20
15
18
8
46
I
40
I
3
10
I
7
3
2
2
2
I
I
3
I
I
I
15
2
2
7
I
I
13
I
I
16
32
12
I
I
Macrosteles quad.
Chlorotettix unicolor
8
2
I
I
Cuema striata
Athysanella attenuata
97
Endria inimica
I
I
I
8
11
18
2
41
15
38
I
2
I
I
I
Dikrella sp.
Amblysellus grex
2
I
Athysanella sinuata
I
Neocolidia tumidifrons
Stenometopiellus cookei
I
2
I
I
I
I
I
2
I
Balclutha punctata
Nesosteles neglectus
Athysanella sp.
2
I '
3
2
Mocuellus caprillus
2
T O T A L (N)
213
1077
384
228
37
6
412
92
2
2451
N U M B E R O F S P E C IE S
17
11
4
13
6
2
10
8
2
28
87
A ppendix A.
1988 Leafhopper Phenology Data.
The Brom us inermis/Medicago sativa Plant Assemblage
LEA FH O PPER
S P E C IE S
Patch 21b
Patch 25b
Patch 26b
Sampling Dates
Sampling Dates
Sampling Dates
6/2/88
Balclutha punctata
Psammotettix lividellus
Macrosteles quadrilineatus
123
89
Aceratagallia spp.
Sorhoanus debilis
40
18
Sorhoanus orientalis
Hebecephalus rostratus
16
Diplocolenus configuratus
2
2
Empoasca sp.
Colladonus geminatus
Endria inimica
Dorycephalus platy.
Doratura stylata
7/20/88 8/25/88 6/2/88
I
5
63
705
823
5
4
29
43
TOTAL
7/17/88 8/22/88 6/16/88 7/17/88 8/22/88
18
4
I
3
135
4
5
18
128
2
37
I
5
87
71
101
31
59
62
2
108
6
8
90
72
7
Latalus missellus
6
Cuema striata
Xerophloea viridis
Dikraneura spp.
Athysanella acuticauda
I
17
2
4
17
20
2
55
106
4
28
I
48
I
I
291
15
303
4
9
41
7
517
1753
135
I
48
366
19
1365
294
2129
97
179
6
9
5
I
1709
672
2677
130
6
2
2
90
2
2
Hecalus spp.
Macrosteles quad.
60
9
10
8
I
10
18
10
3
171
29
23
I
I
Athysanus argentarius
33
7
Chlorotettix unicolor
Elymana sp.
2
5
2
40
7
2
29
Auridius auratus
4
33
Sorhoanus Jlavovirens
8
8
Forcipata Ioca
Commellus sexvitattus
3
I
3
Mocuellus caprillus
Elymana circius
Orocastus labeculus
I
I
I
55
17
72
39
39
Scaphytopius frontalis
2
2
Limotettix sp.
I
I
Texananus sp.
I
Laevicephalus sp.
I
3
3
TOTAL (N)
338
930
851
269
743
45
727
4241
362
8506
NUM BER O F SPECIES
10
8
5
12
11
6
18
22
13
32
88
A ppendix A.
1988 Leafhopper Phenology Data.
The Stipa comata/Bouteloua gracilis Plant Assemblage
LEAFHOPPER
SPECIES
M o c u e llu s c a p rillu s
Patch 7A
Patch IOA
Patch 16A
Sampling Dates
Sampling Dates
Sampling Dates
6/21/91
7/16/91
8/7/91
2
216
24
S o rh o a n u s spp.
89
I
9/13/91
A th y sa n e lla spp.
58
111
28
A c e ra ta g a llia spp.
16
67
823
17
I
4
O r o c a stu s p e r p u s illu s
16
P sa m m o te ttix liv id e llu s
13
E m p o a sc a sp.
5
A u r id iu s o rd in a tu s
4
C o lla d o n u s g e m in a tu s
4
S te n o m e to p ie llu s c o o k e i
3
H e c a lu s spp.
2
D o r y c e p h a lu s p la ty .
I
F le x a m ia fle x u lo s a
6/21/91
7/16/91
8/7/91
18
10
3
79
4
30
9/13/91
175
372
111
7/17/91
8/8/91
l
I
13
117
2
l
418
I
15
17
5
28
I
I
516
I
2
8
I
2
ii
4
I
32
7
3
4
67
5
C h lo r o te ttix u n ic o lo r
4
4
D ik r a n e u ra spp.
I
4
4
2
C u e rn a stria ta
3
3
253
63
11
30
13
16
68
6
68
75
590
8
I
5
H e b e c e p h a lu s ro stra tu s
I
3
I
5
I
C o m m e llu s se x v ita ttu s
I
M a c ro s te le s quad.
6
I
2
3
F le x a m ia a b b re v ia ta
2
I
4
9
4
17
12
I
26
3
2
3
3
R o se n u s c ru c ia tu s
I
I
F r ig a rtu s fr ig id u s
I
I
L im o te ttix sp.
I
I
N e so ste le s n e g le c tu s
D e lto c e p h a lu s va le n s
250
13
8
11
I
6
2
A u r id iu s h e lv u s
428
54
6
4
41
UMBER OF SPECIES
1012
1524
211
398
13
7
275
189
I
O ro c a stu s Ia b ecu Iu s
TOTAL (N)
N
9/16/91
16
39
I
6/18/91
I
I
936
38
547
559
129
32
219
502
706
31
4377
10
7
12
9
8
9
10
6
9
7
27
89
A ppendix A.
1988 Leafhopper Phenology Data.
The Festuca idahoensis/Agropyron spicatum Plant Assemblage
LEAFHOPPER
SPECIES
Patch 21A
Patch 2 SA
Patch 26A
Sampling Dates
Sampling Dates
Sampling Dates
6/27/91
7/18/91
S o rh o a n u s s p p .
298
201
D ik r a n e u r a s p p .
72
I
P sa m m o te ttix liv id e llu s
23
17
16
9
5
5
O ro c o a stu s p e r p u sillu s
C o lla d o n u s g e m in a tu s
A u r id iu s o rd in a tu s
M a c ro s te le s qu a d .
8/6/91
3
I
73
2
36
60
I
M e sa m ia c o lo r a d e n sis
I
I
O r o c a stu s Ia b ecu Iu s
I
D o r y c e p h a lu s p la ty.
I
21
55
41
54
11
9
7
7
2
C h lo r o te ttix u n ic o lo r
D o r a tu r a sty la ta
A u rid iu s a u ra tu s
M o c u e llu s c a p rillu s
5
N e so ste le s n e g le c tu s
6/27/91
7/18/91
417
70
56
12
I
5
6
10
3
2
I
26
2
2
C o m m e llu s se x v ita ttu s
E n d r ia in im ica
11
5
60
2
A th y sa n e lla s p p .
9/16/91
3
5
8/6/91
9/13/91
6/27/91
7/16/91
8/6/91
95
5
13
8
223
6
16
3
16
1309
152
73
I
112
46
49
I
57
I
I
128
98
I
4
I
4
I
12
40
24
I
66
34
20
3
30
4
3
I
38
3
20
2
57
8
2
65
i
7
3
332
164
10
22
20
5
7
123
4
37
2
53
21
93
I
4
4
9
2
43
7
5
I
3
2
I
H e c a lu s s p p .
L im o te ttix s p .
I
I
8
S c a p h y to p iu s s p .
I
I
M a c ro s te le s qu a d .
9/13/91
2
I
I
I
E m p o a sc a s p .
N
2
6
I
I
I
10
I
9
9
I
L a e v ic e p h a lu s s p .
I
X e r o p h lo e a virid is
3
N e o c o lid ia tu m id ifro n s
I
3
6
12
6
12
I
C u e rn a stria ta
2
D ip lo c o le n u s co n fig .
2
4
I
4
7
I
B a lc lu th a p u n c ta ta
■:'■
I
F o rc ip a ta Ioca
I
I
I
F r ig a rtu s fr ig id u s
..
E ly m a n a c irciu s
18
18
A u rid iu s h e Ivus
I
I
I
L a ta lu s m issellu s
I
P a r a p h le p s iu s s p .
TOTAL (N) 454
NUMBER OF SPECIES
I
12
I
547
92
120
518
257
72
106
274
309
121
6
2876
21
13
7
11
14
9
9
11
19
9
4
35
90
A ppendix A.
1988 Leafhopper Phenology Data.
The Agropyron cristatum/Medicago sativa Plant Assemblage
LEA FHO PPER
SPECIES
6/21/91
S o rh o a n u s spp.
26
P sa m m o te ttix livid ellu s
15
Patch 7 b
Patch 16b
Patch 17b
Sampling Dates
Sampling Dates
Sampling Dates
7/16/91
A c e ra ta g a llia s p p .
13
79
D ik r a n e u r a s p p .
11
H e c a lu s s p p .
6
6
5
3
2
I
3
4
M o c u e llu s c a p rillu s
O r o c a stu s p e r p u sillu s
M a c ro s te le s q u a d rilin e a tu s
C o lla d o n u s g e m in a tu s
R o se n u s c ru c ia tu s
6
806
9/16/91
6/18/91
2
14
29
6
6
I
7/17/91
8/8/91
N
9/16/91
6/18/91
7/17/91
8/8/91
9/16/91
3
39
I
25
31
2
4
17
6
42
138
6
4
6
I
I
43
5
I
8
1120
6
22
146
8
38
56
6
I
I
I
2
7
8
I
2
17
2
125
A th y s a n e lla s p p .
8/7/91
D o r y c e p h a lu s p la ty .
6
O ro c a stu s la b e c u lu s
2
C o m m e llu s se x v ita ttu s
I
4
5
237
63
35
19
5
106
I
548
3
6
2
58
I
I
I
C u e rn a stria ta
S te n o m e to p ie llu s c o o k e i
I
I
I
H e b e c e p h a lu s ro stra tu s
I
I
5
3
D o r a tu r a sty la ta
H a r d y a d e n ta ta
5
I
4
2
M a c ro s te le s qu a d .
3
2
I
A m b ly se llu s g r e x
I
TOTAL (N)
88
221
819
11
57
326
249
22
56
138
68
11
2066
NUMBER O F SPECIES
10
8
6
4
7
6
7
5
5
7
5
4
20
91
A ppendix A.
1988 Leafhopper Phenology Data.
The Brom us inermis/Medicago sativa P lant Assemblage
LEA FHO PPER
SPECIES
D ik r a n e u r a s p p .
C o lla d o n u s g e m in a tu s
6/27/91
Patch 25b
Patch 26b
Sampling Dates
Sampling Dates
7/18/91
8/6/91
89
78
P sa m m o te ttix livid ellu s
35
13
S o rh o a n u s s p p .
Patch 21b
Sampling Dates
M a c ro s te le s q u a d rilin e a tu s
4
D ip lo c o le n u s c o n fig u ra tu s
3
A th y s a n e lla s p p .
2
A m b ly se llu s g r e x
I
A c e ra ta g a llia s p p .
E n d ria in im ica
D o r a tu r a sty la ta
I
I
13
8
I
2
92
18
16
449
2
10
N
9/16/91
6/27/91
7/16/91
8/6/91
9/16/91
6/27/91
7/16/91
8/6/91
9/13/91
16
7
107
3
35
8
25
4
10
55
2
72
5
32
2
12
6
23
8
14
6
52
48
33
3
58
19
3
9
65
i
I
11
103
94
86
57
48
2
68
112
12
64
I
19
108
I
11
128
1682
2576
42
I
17
68
4
4
18
123
7
36
2933
4278
78
1158
1983
203
147
28
188
71
14
5833
I
8943
963
I
I
N e s o s te le s n e g le c tu s
332
431
I
D o r y c e p h a lu s p la ty.
426
107
C o m m e llu s se x v ita ttu s
2
A th y s a n u s a rg e n ta riu s
8
16
18
L a ta lu s m issellu s
I
A u r id iu s o rd in a tu s
7
3
4
10
I
13
5
53
42
33
25
112
32
13
52
95
E ly m a n a c irciu s
6
28
17
51
H e c a lu s s p p .
3
20
6
29
M a c ro s te le s quad.
2
6
3
24
13
C h lo r o te ttix u n ic o lo r
4
4
O ro c a stu s la b e c u lu s
I
I
7
8
3
L im o te ttix s p .
S c a p h y to p iu s s p .
2
4
I
B a lc lu th a p u n c ta ta
11
8
6
C u e rn a stria ta
I
M e sa m ia c o lo r a d e n sis
I
I
I
116
18011
8
27
TOTAL (N)
NUMBER O F SPECIES
225
142
473
205
272
4622
214
174
8
7
7
5
10
16
13
9
92
328
8
7770 3470
19
18
A PPEN D IX B: T H E LEA EH O PPER SPECIES C O LLEC TED D U RIN G TH IS STUDY
93
Appendix B. The Leafhopper Species Collected during this Study.
subfamily AGALLDNAE
subfamily DELTOCEPHALINAE cont'd
Aceratagallia sanguinolenta Provancher
Aceratagallia uhleri (Van Duzee)
genus Scaphytopius Ball
Sorhoanus debilis (Uhler)
Sorhoanus flavovirens (Gillette and Baker)
Sorhoanus orientalis (Delong and Davidson)
Stenometopielus cookei (Gillette)
Streptanus conjinis (Rent.)
genus Texananus Ball
subfamily BALCLUTHINAE
Nesosteles neglectus (Delong and Davidson)
Balcluthapunctata (Thunb.)
subfamily DELTOCEPHALINAE
Am blysellusgrex (Oman)
subfamily DORYCEPHALINAE
Athysanella (Amphipyga) acuticauda Baker
Dorycephalus platyrhynchus Osborn
Athysanella (Amphipyga) attenuata Baker
Athysanella (Amphipyga) occidentalis Baker
subfamily HECALEfAE
Athysanella (Athysanella) robusta (Osborn)
Hecalus major (Osborn)
Athysanella (Gladioneura) sinuata (Osborn)
H ecalusviridis (Uhler)
Athysanella (Athysanella) terebrans (Gillette and Baker)
Athysanella (Athysanella) utahna (Osbom)
subfamily IASSEfAE
Athysanus argentarius M etcalf
Prairiana cinerea (Uhler)
Auridius auratus (Gillette and Baker)
Prairiana subta Ball
Auridius helvus (Delong)
subfamily LEDREfAE
Auridius ordinatus (Ball)
Xerophloea viridis (Fabricius)
Chlorotettix unicolor (Fitch)
Cicadula quinquinotdta (Boh.)
subfamily NEOCOLIDIEfAE
Colladonus geminatus (Van Duzee)
Neocolidia tumidifrons Gillette and Baker
Colladonus montanus (Van Duzee)
Commellus semicolon Hamilton
subfamily TETTIGELLEfAE
Commellus sexvitattus (Van Duzee)
Cuem a striata (Walker)
Deltocephalus valens Beamer and Tuthill
Diplocolenus configuratus (Uhler)
subfamily TYPHLOCYBEfAE
D oraturastylata (Boh.)
Dikraneura shqshone Delong and Caldwell
Elymana circius Hamilton
g em s Dikrella Oman
Endria inimica (Say)
genus Empoasca Walsh
Endria rotunda (Beamer)
Forcipata Ioca (Delong and Caldwell)
Flexamia abbreviata (Crumb)
Flexamia flexulosa (Ball)
Frigartusfrigidus (Ball)
Hardya dentata (Osborn and Ball)
Hebecephalus rostratus Beamer and Tuthill
Idiodonus aurantiacus (Provancher)
g e m s Laevicephalus Delong
Latalusm isellus (Ball)
g em s Lonatura Osbom and Ball
M acrosteles quadrilineatus (Forbes)
Mesamia coloradensis Gillette and Baker
M ocuellus caprillus Ross and Hamilton
Norvelina seminuda (Say)
Orocastus labeculus (Delong)
Orocastus perpusillus (Ball and Delong)
Paraphlepsius occidentalis (Baker)
Pinumius areatus (Stal)
Psammotettix lividellus (Zettinger)
Rosenus cruciatus (Osborn and Ball)
94