Vol. 24 No. 1 - Michigan Entomological Society

Transcription

Spring 1991 Vol. 24, No.1
THE GREAT LAKES ENTOMOLOGIST PUBLISHED BY
THE MICHIGAN ENTOMOLOGICAL SOCIETY THE GREAT LAKES ENTOMOLOGIST
Published by the Michigan Entomological Society
No. 1
Volume 24
ISSN 0090-0222
TABLE OF CONTENTS
Light trap records of Phyllophaga (Coleoptera: Scarabaeidae) in Wisconsin,
1984-1987
Robert A. Dahl and Daniel L. Mahr ......................................... .
Acoustic signals of Graminella nigrifrons (Homoptera: Cicadellidae)
S. E. Heady and L. R. Nault ................................................ 9 No intersexual differences in host size and species usage in Spa/angia endius
(Hymenoptera: Pteromalidae)
B. H. King ............................................................... 17 Acrobasis shoot moth (Lepidoptera: Pyralidae) infestation-tree height link in a
young black walnut population
George Rink, Barbara C. Weber, D. Michael Baines and David T. Funk .......... 21
Host plant suitability and a test of the feeding specialization hypothesis using
Papilio cresphontes (Lepidoptera: Papilonidae)
J. Mark Scriber and Robert V. Dowell ....................................... 27 Geographic distribution of Siphonaptera collected from small mammals on
Lake Michigan Islands
William C. Scharf ......................................................... 39
Acorns as breeding sites for Chymomyza amoena (Diptera: Drosophilidae) in
Virginia and Michigan
Henretta Trent Band ....................................................... 45
List of the Lepidoptera of Black Sturgeon Lake, Northwestern Ontario, and dates
of adult occurrence
C. J. Sanders ............................................................. 51 COVER ILLUSTRATION
A mourning cloak butterfly, Nympha/is antiopa (L.)
at Ludington State Park, ML
Photograph by M. F. O'Brien
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Copyright
1991. The Michigan EntomOlogical Society
1991
THE GREAT LAKES ENTOMOLOGIST
LIGHT TRAP RECORDS OF PHYLLOPHAGA (COLEOPTERA:
SCARABAEIDAE) IN WISCONSIN, 1984-1987
Robert A. Dahl and Daniel L. Mahr l
ABSTRACT
Phyllophaga adults (June beetles) were surveyed from 1984 through 1987 at five
locations in Wisconsin using black light traps. Data were collected at each location
for three consecutive years. A total of 9,259 specimens representing 13 species were
collected during the survey. Species captured, sex ratios, and flight periods for
abundant species were recorded for each location. Species abundance differed from
previous surveys of Phyllophaga in Wisconsin. Possible reasons for observed shifts
in species abundance are discussed.
Phyllophaga larvae (white grubs) feed upon the roots of turfgrasses and other
desirable plants, whereas adults (June beetles) feed on the foliage of trees and
shrubs. In Wisconsin, most Phyllophaga are thought to have a three year life cycle,
spending most of it as larvae in the soil (Luginbill and Painter 1953). Much is known
about adult host plant preferences (Forbes 1916, Travis 1933, Ritcher 1940, Cham
berlin et al. 1943), but although Ritcher (1940) made some general observations on
white grub hosts in Kentucky, the host relationships of Phyllophaga larvae are
poorly understood.
Since the last study of Wisconsin Phyllophaga (Chamberlin et al. 1943), agricul
tural practices and land use patterns have changed considerably. Major differ
ences include increased use of persistent and non-persistent soil insecticides, modi
fied weed control practices, increased irrigation in some areas, adoption of crop
rotation, shifts in pasture forage species, adjustments in fertilization practices,
and escalating suburbanization. These and other alterations in land use could
seriously impact the species composition of the soil insect fauna, including white
grubs.
Changes in the species complex of Phyllophaga within a region may be a reflec
tion of modifications in larval and/or adult habitats. Proximity to adult host plants
and soil moisture are two major factors influencing Phyllophaga populations
(Forbes 1916, Sweetman 1931).
Light traps are commonly used to surveyor monitor Phyllophaga adults (Sanders
and Fracker 1916, Forbes 1916, Travis 1933, Henry and Heit 1940, Neiswander
1963, Lim et aL 1979). This paper reports a light trap study in Wisconsin which
surveyed the species of Phyllophaga and their relative abundance at five different
locations in the state. Goals of the study were to; (l) examine for species shifts since
earlier studies and (2) analyze for possible periodicity in adult emergence. Addition
ally, we review the brood theory concept as it relates to Phyllophaga yearly abun
dance.
1Department of Entomology, University of Wisconsin, 1630 Linden Drive, Madison, WI
53706.
2
VoL 24, No. I
•
GOLF COURSE
•
AGRICULTURAL
Figure 1. Location of blackIight traps for surveying Phyllophaga spp. adults in Wisconsin
(1984-87).
MATERIALS AND METHODS
Four-baffled blacklight traps (Ellisco Inc. cat. no. 110103-2) equipped with ultra
violet fluorescent bulbs (18" Norelco 15 watt bulb FI5 T8/BL) were used to monitor
Phyllophaga adults at five locations in Wisconsin from 1984 to 1987. Three consecu
tive years of trapping were conducted at each location. From 1984 through 1986,
single traps were operated continuously from mid-April to mid-July on each of three
suburban golf courses in Madison, Brookfield, and Menasha (Figure 1). From 1985
through 1987, two traps were operated in Mazomanie and Lancaster, agricultural
locations in the southern part of the state. A dichlorvos-impregnated strip in the
collecting pan of each trap killed the trapped insects.
Trap catches were collected weekly, placed in labeled paper bags and stored in a
refrigerator until processed. Phyllophaga were sorted according to species, sexed and
counted. Identification of species was according to the keys of Luginbill and Painter
(1953). Voucher specimens from this study are deposited in the Insect Research Collec
tion, Department of Entomology, University of Wisconsin, Madison.
1991
3
RESULTS AND DISCUSSION
Species Collected
Thirteen species and 9,259 specimens of Phyllophaga were collected during the
four years of the study (Table 1). All 13 species had been previously recorded for
Wisconsin although previous studies recorded greater diversity of Phyllophaga than
our study. For example, Sanders and Fracker (1916) collected 17 species at five
Wisconsin collection sites with light traps whereas Chamberlin et aJ. (1943) collected
19 species at 39 different locations in Wisconsin by handpicking beetles from host
plants at night. Data from Iowa (Travis 1933) and northern Illinois (Forbes 1916)
also showed greater diversity of Phyllophaga than our study. The lower species
diversity observed in our study may have been due in part to the restricted number of
habitats associated with our survey locations.
The most commonly collected species in our study was P. JutiUs LeConte, which
accounted for 710,70 of the total. The Madison sites surveyed previously (Sanders and
Fracker 1916) and during our study show differences in species abundance. P.
rugosa Melsheimer and P. Jusca Froelich comprised 990,70 of the June beetles col
lected by Sanders and Fracker (1916) at Madison but in our study, P. JutiUs and P.
anxia LeConte collectively accounted for 870,70 of the June beetles captured in Madi
son. Conversely, P.rugosa and P. Jusca represented only 5.9070 of our total and P.
JutiUs and P. anxia constituted < 1% of the total caught by Sanders and Fracker
(1916). Several reasons may explain the difference in species abundance between our
study and that of Sanders and Fracker (1916). First, it is highly probable that our
light trap locations within Madison differed from those of the 1916 study. Second,
the type of adult host trees and their proximity to light traps may have influenced
the array of Phyllophaga species caught (e.g. Forbes 1916). Most importantly, sig
nificant changes in land use patterns have occurred in Madison over the 70 years
between studies and it is likely that habitats of larvae and/or adults were altered
during this time. Our Madison location was on a golf course adjacent to the Univer
sity of Wisconsin Arboretum, a vegetation ally diverse area where many native Wis
consin plant communities are recreated on one tract of land. However, data from a
light trap on another golf course in the Madison area showed the same two most
abundant Phyllophaga species (P. Jutilis and P. anxia) for a two year period (1985
and 1986) (unpublished). The most abundant species at each location in our study,
except Mazomanie, was P. Jutilis (Table 1). The most abundant species at the
Mazomanie location was P. implicita Horn. Each location had a different species
complex and only three species were common to all five trap locations. Shenefelt
and Simkover (1951) found that the Phyllophaga species composition differed
greatly between localities and concluded that these differences were a manifestation
of the environmental conditions under which the insects exist (e.g. edaphic factors).
Sex Ratios of Phyllophaga Taken at Light Traps
Sex ratios for the most abundant species in the present study, given as the fraction
of males, are as follows: P. anxia 0.88 (n = 75); P. crenulata Froelich 0.97 (n = 62); P.
Jusca 0.76 (n "" 71); P. JutiUs 0.94 (n = 6214); P. gracilis Burmeister 0.75 (n = 16); P.
hirticula 0.89 (n=28); P. implicita 0.52 (n 1423); P. rugosa 0.90 (n=919). Gener
ally, light traps capture more male Phyllophaga than females (e.g. Travis 1933, Lim
et aL 1979) although some studies report differing results concerning sex ratios for
certain species. For example, in three light trap studies involving P. anxia. Morofsky
(1933) collected more females than males while Neiswander (1963) and Sanders and
Fracker (1916) collected more males than females. In our study P. anxia captures
were male biased. Only P. implicita had a nearly equal sex ratio and reasons for this
were not apparent. Several factors could contribute to differences in sex ratios using
light traps. For example, differences in flight habits and differential phototaxis
(wavelength of light used in trap) among the sexes are two factors which could affect
sex ratios.
""" Table I. Phyllophaga spp. beetles collected at light traps in Wisconsin, 1984-1987.
Phyllophaga
BROOKFIELD
species
-~
0
0
0
0
0
0
200
140
130
0
0
0
total
- -....... - - . -.......
~-.-
1986
0
0
0
0
0
0
0
4
0
0
I
1985
129
9
1
I
0
119
8
I
0
I
0
I
0
0
0
0
0
0
196
MENASHA
LANCASTER
MAZOMANIE
i
-
~-
1984
jutilis
rugosa
anxia
jusca
implicita
crenulata
tristis
hirticula
balia
gracilis
in versa
jorsteri
prunina
MADISON
-J
0
'---------.-...- -...... -~
~~
1984
98
0
9
5
1
14
1
0
2
0
0
0
0
130
~-
.......
-
1985
1986
306
27
20
0
5
4
2
0
0
0
0
0
I
30
1
22
0
0
6
1
365
~-
1985
1986
1985
1046
30
7
0
0
0
0
0
0
0
0
0
0
345
0
0
0
0
0
1984
487
3
0
0
0
0
0
0
0
0
0
0
0
358
79
11
8
103
8
0
0
1
2
0
I
0
60
490
1083
387
0
-~
~~
4()
1
0
I
0
0
0
0
0
0
0
0
571
-~
1985
1291
640
0
26
15
0
0
1986
3
1
2
0
271
11
1987
0
9
1
3
977
15
0
0
0
0
0
0
0
0
0
0
14
I
0
0
10
288
1020
1986
220
2
0
1987
0
0
1
0
0
5
0
0
0
0
0
0
0
0
1944
145
0
25
44
4
0
18
0
0
0
0
0
1983
232
2180
5
i
I
x:tI1
o
~
~
~
tI1
Ul
tI1
Z
d~
o
r
I
o
o.....
Ul
-J
i
~
Z
o
1991
5
200
Brookfield
150
"'0
CD
100
:::J
0
RS
50
I
0
u 400
.(1) 300
..::::: 200
100
~
0
120
90
60
Q" 30
0
::::::
0
... ---_ ......
/
/
.....
0
I
CD
.Q
E
200
:::J
Z
100
,
,,
"co. __
Menasha
-----', ,
/
/
~ "","
~
/
,-- ...
=-*-
...
~
--- .......
/
/
...
/'
-"
.../~
;' "
...
...
....
1984
1985 1986
----
Lancaster
'-,
'
"
__L ..........>.,
'-, '--
,/' , ,
/
/'
/'
-...
I
--
Madison
/
./
1984
1985
. ,,--. 1986
--
/
.2
1200
Q"
-
,
1984
1985
1986
. " .... """ . '" ~
/
~
.a::"
900
600
300
0
--- -
,
...
1985
1986
1987
Mazomanie
1985
1986
1987
0
A{\~ r:,1"'- r:,/\\ r:,/\\\ r:,/\~ to/\ to/\\ to/\\\ to/\~
Month/Week
Figure 2. Number of Phyllophaga jutilis beetles caught per week at light trap locations in
Wisconsin, 1985-1987.
Seasonal Flight Periods and Yearly Trap Results
The three most abundant Phyllophaga species in our study were P. jutilis, P.
impiicita, and P. rugosa. Other Phyllophaga species (Table 1) were not well repre
sented and are not discussed. The three most abundant species have similar adult
seasonal activity periods. The fourth week of April through the end of June brackets
the period of highest activity of adult Phyllophaga species. Many environmental
factors (such as soil moisture and temperature, air temperature, and local rain fall
patterns) potentially influence developmental rate or flight activity. Such factors
could result in different flight periods for a species in areas of close proximity. For
example, in 1985 Mazomanie and Lancaster had similar flight activity periods for P.
jutilis (Fig. 2) while the activity periods for P. implicita in 1987 at the same two
locations were quite different (Fig. 3).
Peak captures of P. jutilis in 1984 occurred during the same time period for the
three golf course locations (Fig. 2). In 1985 all trap locations had earlier flight
activity of P. jutilis than other years; the two agricultural locations had earlier peak
flights than the three golf course sites (Fig. 2). Flight activity of P. jutilis in 1986 was
6
"0
•
20
L
~
+-
Q.
15
"u 10
.... .:!
o'u
:::::
5
;~
0
..Q ' It
300
Z
200
E
::::JO)
It
.t::
Q.
,,2
100
if
0
?
Vol. 24, No.1
Lancaster
1985
1986
1987
/"................
;",
;'
/
Mazomanie
,
1985
"
/.;:.. ......::;...... -:..:.::::.:...... ~.~:~~~:::.:.... ""
1986
1987
A,/\'J c:.,1\ c:.,1\\ c:.,1\\\ c:.,1\'J fl/\ fl/\\ fl/\\\ fl/\'J
Month/Week
Figure 3. Number of Phy/lophaga implicita beetles caught per week at light trap locations in
more variable temporally than the previous two years. Both agricultural locations
had low numbers of P. julilis captured in 1986. Total P. julilis collected at all sites
was lowest in 1986, except at Mazomanie (Table 1). In 1987, Lancaster had the
largest capture of P. jUliUs for any year or location, whereas Mazomanie had no P.
jutilis captured in 1987, the lowest for any year at any location during the study.
Abundance of P. rugosa was greatest in 1985 at all sites except Menasha (Table 1).
Captures were very small in 1984 and 1986 at all locations, Menasha excluded.
Captures differed substantially in 1987 between Mazomanie and Lancaster. The
total capture of P. rugosa at Lancaster in 1987 was substantially greater than at
Mazomanie (Fig. 4).
Seasonal flight periods were similar for P. rugosa at Brookfield and Menasha in
1984 (Fig. 4). In 1985, seasonal flight periods were earlier than 1984 (for Brookfield
and Menasha) and similar for all locations except Mazomanie which peaked later
(Fig. 4). Peak flight periods in 1986 were similar among the golf course locations
and 1987 peak flight periods were similar between the two agricultural locations
(Fig. 4).
P. implicita was collected at all five sites but was well represented only at the two
agricultural locations (Table 1). More P. implicita were captured yearly at Mazoma
nie than at Lancaster, and Mazomanie trap catches increased with each year of
trapping (Table 1). Peak captures of P. implicita were generally earlier at Mazoma
nie for a given year (Fig. 3). Although P. implicita was captured mainly at the
agricultural locations, the total number captured represented> 150/0 of the 9,259
beetles collected.
The Brood Theory in Phyllophaga
Several authors have alluded to a three year synchronous life-cycle ("broods") for
Phyllophaga (Neiswander 1963, Hammond 1954, Ritcher 1940). The idea of syn
chronous and periodic emergence of June beetles may have originated from Euro
pean work. For example. Forbes (1916) briefly mentioned a European origin for
forecasting white grub outbreaks but lists no references. Unfortunately the notion of
synchronicity with respect to Phyllophaga populations has been perpetuated without
sound data. Neiswander (1963) claimed that entomologists have "essentially
1991
6
"...
-
,---------- A'-~...;,.··_--=.::;.:.:.:..j:::
2
:::J
Q. 11$
Brookfield
4
I\)
0
/\
(J
til
20
0
10
'"
0
15
10
5
I/)
~
&
til
.c::
:::::~
..
I\)
..Il
E
:::J
Z
!
Menasha
/ " , / \.
,
",
"
/'
.:'
"'\
'" / ...... ~.:
:..,
""
. ... ___ ---.
,. ~
1984
--- 1985
...... 1986
..
~~~,
Madison
1984
- -- 1985
...... 1986
---
0
Lancaster
~200
....0 ...
7
,
100 /
0
40
30
20
10 0
• 1985
...... 1986
_.- 1987
/
.-.-.-~-'----------~;;.:--"-';;":'-::;-.;:-.
/
/
/
'
Mazomanie
,,
,;.:.:::.:..: _____ • __ ~:.:..:._.
/J.1\<oJ
'fI1\
'fI1\\ 'fI1\\\ 'fI1\<oJ
,
1985
...... 1986
' _ . - 1987
6/\ 6/\\ 6/\\\ 6 / \<oJ Month/Week
Figure 4. Number of Phyllophaga rugosa beetles caught per week at light trap locations in
accepted the early alignment of broods without subsequent study of populations."
The life cycles of Phyllophaga species generally exhibit a periodicity of two to four
years but individuals of the same species may differ greatly (e.g. Davis 1916).
Determination of life cycle length in Phyllophaga is complex; the interplay of tern·
perature, food quality and quantity, soil conditions and larval population density
may all be involved. In particular, cumulative seasonal soil temperatures can have a
strong influence on developmental time. For instance, Davis (1916) found P. drakii
Kirby to have a three year life cycle at Lafayette, Indiana and a four year life cycle at
Trout Lake, in northern Wisconsin, Furthermore, he pointed out that the combined
seasonal temperatures in Lafayette for three years were approximately the same as
for four years at Trout Lake. Additionally, Shenefelt and Simkover (1951) found P.
tristis Fabricius to have a two year life cycle in central Wisconsin but a one year life
cycle at the same latitude in Michigan. Michigan's more moderate climate may
account for the shorter life cycle. The roles of food quality and larval density are
more difficult to assess. These factors could, separately or together, account for
differences in life cycle length for individuals of the same species in the same
location. Lloyd and White (1976) proposed that Magicicada species might emerge
four years early with crowding of early instar nymphs. These factors make predic
tion of destructive broods difficult except on a very local basis.
Mahr (1984) predicted that 1986 would be a major flight year for P. rugosa in
Wisconsin. No trap location in this study supported that prediction. Although the
Menasha location had the largest capture of P. rugosa in 1986, the total was fairly
insignificant and P. rugosa was not the most abundant species in any year. The
literature shows discrepancies in major flight years and relative abundance for a given
8
Vol. 24, No. 1
species in different areas (Forbes 1916, Sanders and Fracker 1916). It is therefore
difficult, if not impossible, to compare results of different studies in predicting Phy
l/ophaga outbreaks, even in the same locale. In some areas a shift in the most abun
dant brood can occur over several years (Neiswander 1963). The supposition that most
June beetles have synchronous life cycles appears to be a dubious generality on a
regional basis. In restricted geographical areas populations of Phyl/ophaga may be
temporarily synchronous due to localized environmental conditions. However, it is
likely that this synchronicity breaks down through time and the concept should not be
extrapolated on a larger geographical or temporal scale.
ACKNOWLEDGMENTS
Funding for this study was provided, in part, by the Wisconsin Turfgrass Associa
tion and the University of Wisconsin College of Agricultural and Life Sciences, and
by Hatch Grant nos. 2011 and 2566. We thank Ed Arnold of the Wisconsin Dept. of
Agriculture, Trade and Consumer Protection and the following golf course superin
tendents: Randall Smith, Nakoma Country Club, Madison; Roger Bell, Northshore
Golf Club, Menasha; Jerry Kushasky, Westmoor Country Club, Brookfield. We
also thank Phil Pellitteri for assistance in identification of many June beetles.
LITERATURE CITED
Chamberlin, T. R., C. L. Fluke and J. A. Callenbach. 1943. Species, distribution, flight and
the host preferences of June beetles in Wisconsin. J. Eeon. Entomol. 36:674~680.
Davis, J. J. 1916. A progress report on white grub investigations. J. Econ. Entomol.
9:261-281.
Forbes, S. A. 1916. A general survey of the May-beetles (Phyllophaga) of Illinois. Ill. Agric.
Exp. Sta. Bull. 186:215-257.
Hammond, G. H. 1954. Long-term fluctuations in populations of white grubs (Phyllophaga
spp.) in sod in Eastern Ontario. Empire J. Exp. Agr. 22:59-64.
Henry, H. K. and C. E. Heit. 1940. Flight records of Phyllophaga (Coleoptera: Scarabaeidae).
Entomol. News. 51:279~282.
Lim, K. P., K. M. Toohey, W. N. Yule and R. K. Stewart. 1979. A monitoring program for the
common June beetle, Phyllophaga anxia (Coleoptera: Scarabaeidae), in southern Quebec.
Can. Entomo!. III: 1381-1397.
Lloyd, M. and J. A. White. 1976. Sympatry of periodical cicada broods and the hypothetical
four-year acceleration. Evolution 30:786-801.
Luginbill, P. and H. R. Painter. 1953. May beetles of the United States and Canada. U. S.
Dept. Agric. Tech. Bull. 1060. 102 p.
Mahr, D. L. 1984. Turfgrass disorder: White grubs. Univ. Wisc. Ext. Pub!. A3275. 3 p.
Morofsky, W. F. 1933. Distribution of May beetles (Phyllophaga) in Michigan. J. Econ.
EntomoL 26:831~834.
Neiswander, C. R. 1963. The distribution and abundance of May beetles in Ohio. Ohio Agric.
Exp. Sta. Res. Bull. 951. 35p.
Ritcher, P. O. 1940. Kentucky white grubs. Ky. Agric. Exp. Sta. Bull. 401:1-151.
Sanders, J. G. and S. B. Fracker. 1916. Lachnosterna records in Wisconsin. J. Econ. Entomol.
9:253-261.
Shenefelt, R. D. and H. G. Simkover. 1951. Notes on habits and "broods" of June beetles.
Entomol. News. 57:219-223.
Sweetman, H. L. 1931. Preliminary report on the physical ecology of certain Phyllophaga
(Coleoptera: Scarabaeidae). Ecology 12:401-422.
Travis, B. V. 1933. Notes on the habits of June beetles in Iowa (Phyllophaga-Coleoptera).
Iowa State CoiL J. Sci. 7:397-406.
1991
9
ACOUSTIC SIGNALS OF GRAM/NELLA NIGRIFRONS
(HOMOPTERA: CICADELLIDAE)
S. E. Heady and L. R. Nault I
ABSTRACT
The deltocephaline leafhopper, Graminella nigrifrons, produces low intensity sub
strate transmitted vibrations (signals) to facilitate location of virgin females by
males during courtship. In the laboratory, signals produced on maize leaves were
received by a phonographic cartridge, amplified, and analyzed on an oscillograph
and sonograph. Male calls, that are produced spontaneously, are complex, consist
ing of three consecutive sections. Section 1 consists of ca. 3 sec of irregular clicks.
Section 2 has ca. 4 sec of repeated phrases consisting of a continuous series of 0.4 sec
chirps and a roll. Section 3 consists of ca. 5 sec of an intermittent series of 0.2 sec
chirps and a roll. Female calls are produced in response to male calls. Female calls
are simple compared to male calls and consist of ca. 4-5 sec of low frequency
clicking. Signal patterns of G. nigrifrons are compared to those of other leafhoppers
and evolutionary scenarios are presented to account for the observed gender differ
ences in signals.
The black faced leafhopper, Graminella nigrifrons (Forbes), is one of the most
common leafhoppers occurring in grasslands of the Eastern United States (Kramer
1967). G. nigrifrons is the primary vector of the virus causing one of the most
important stunting diseases of maize (Zea mays mays) in the United States. The
pathogen, initially described as the 'Ohio corn stunt agent' by Rosenkranz (1969).
was later identified as maize chlorotic dwarf virus (MCDV) (Gingery 1988 and
references therein). In the corn belt states, MCDV has been reported in Ohio,
Illinois, and Indiana, but is generally a problem only in areas where its alternate
host, johnsongrass (Sorghum halepense L.), occurs (Gordon et al. 1981). Use of
tolerant maize genotypes can reduce losses in some years and locations. MCDV
remains the most destructive and widespread maize virus in the Southeastern and
adjacent maize growing regions of the United States (Gordon and Nault 1977).
The biology of G. nigrifrons and its role as the vector of MCDV are well docu
mented. Studies have been conducted on field biology (Boyd and Pitre 1968, Stoner
and Gustin 1967), host range (Boyd and Pitre 1969), host utilization (Hunt and
Nault 1990, Larsen et aI. 1990). distribution (Douglas et al. 1966, Durant 1968,
Durant and Hepner 1968), and vector relationships (Nault et al. 1973, Choudhury
and Rosenkranz 1983, Knoke et al. 1983). The reproductive biology of G. nigrifrons
has been studied in terms of laboratory life tables (Sedlecek et al. 1986); however,
studies on the mating behavior were lacking. Leafhoppers were discovered to pro
duce low intensity substrate signals by Ossiannilsson (1949). These signals are inte
gral in courtship and facilitate males locating virgin females for copulation. To aid
IDepartment of Entomology, Ohio Agricultural Research and DevelopmentCenter,The
Ohio State University, Wooster, OH 44691.
10
Vol. 24, No. 1
our understanding of the mating behavior of G. nigrijrons, male and female signals
are described herein. Other aspects of the mating behavior, particularly those that
may influence vector movement and disease spread, were conducted concurrently
(Hunt 1988).
Acoustic recordings were made using adults taken from laboratory colonies kept
in rearing cages (D'Arcy and Nault 1982) at 26-28°C with a photoperiod of 12:12
(L:D). Late-instar nymphs were individually separated to obtain virgin adults for
recording (Heady et al. 1986). To record signals, a leafhopper was placed on a maize
leaf piece (ca. 15 cm length) and quickly covered with a plastic dome (l cm diam)
(Heady 1987). The leaf piece was laid over a phonographic cartridge (Electro Voice
5146) so that it lightly touched the needle. Signal output from the cartridge was sent
to a preamplifier (Omega EQ-25), a DC amplifier at l00x (Dana 3640), displayed on
an oscilloscope (Dumont/Fairchild 766), monitored with earphones, and recorded
on a tape recorder (Nagra E) using magnetic tape (3M 250) set at 19 cm/s. Record
ings were made in a laboratory where temperatures ranged from 25 ± 2°C. Calls on
acoustic tape were printed using a pen and ink polygraph (Grass Model 7, D.C.
driver amplifier)(Heady et al. 1986). For male G. nigrijrons, the average duration of
a spontaneous call was measured from 30 randomly chosen individuals. Characteri
zations of portions of the male signals were analyzed from oscillograms of five calls
per individual and 10 individuals using a repeated measures analysis of covariance,
with temperature as the covariable. Calls of female G. nigrijrons were analyzed
from eleven individuals. Frequency spectra were printed using a DSP Sona-Graph
(Model 5500, Kay Elemetrics Corp.).
Call terminology is described based on onomatopoeic interpretation and by pulse
structure (Alexander 1967, Booij 1982, Heady et al. 1986, Huber et al. 1989).
Leafhopper call repertoire includes male calling signals and male and female court
ship signals. These calls are made up of sections composed of phrases, which are
themselves made up of chirps, dicks, rolls, and intervals of silence. Chirps are
distinct sounds to the human ear and are composed of simple or complex pulses of
sound. Clicks have shorter durations than chirps. A roll is composed of a chirp and
3-6 pulses.
RESULTS
G. nigrijrons males spontaneously produced calling signals with mean durations
of 15.2 (S.E.
1.4, N = 30) sec. In the presence of a responding female, the male
would call almost continuously as he walked to the female and joined genitalia. The
male calling signal of G. nigrijrons was composed of three sections (Fig. lA).
Section 1 consisted of irregular clicks which were produced with increasing fre
quency over ca. 3 sec. Section 2 consisted of ca. 4 sec of repeated phrases consisting
of a continuous series of 5-9 chirps and a roll (Fig. IB). Section 3 has ca. 5 sec of
repeated phrases consisting of an intermittent series of 1-4 chirps and a roll (Fig.
IC). The repeating phrase in section 2 was longer than the repeating phrase in
section 3, 0.4 and 0.2 sec, respectively (Table 1). The rolls found in section 2 and 3
were not significantly different in duration and pulse rate (Table 1). Additionally,
the rate of roll production (rolls/msec) in section 2 and 3 was not significantly
different (P >0.20). Thus the main difference between section 2 and 3 was the
chirping before rolls. When the chirps, rolls, and silent interval between phrases
were totaled (Table 1, phrase duration + phrase interval) and compared for section
2 and 3, section 3"'phrases (x = 597.8, S.E.
9.9) were only slightly longer than
17.2). Temperature did not significantly (P>
section 2 phrases (x == 537.5, S.E.
1991
Table I. -
11
Male calling signal variables of Graminefla nigrijrons.
Section 2a
Phrase Duration (msec)
Phrase Interval (msec)
Chirp Rate (chirps/sec)b
Roll Duration (msec)
Roll Pulse Rate (puI es/sec)b
Roll Rate (rolls/sec) 6
II:
449.4 (15.7)
88.1 (10.2)
0.02 (0.0003)
64.6 (1.1)
0.15 (0.0044)
2.00 (0.043)
(S.E.) Section 3a 215.8 (4.5)
382.0 (10.6)
64.6 (0.9)
0.15 (0.0066)
1.87 (0.064)
to
1.
b Values are temperature corrected for 24°C.
0.20) affect duration of phrases or intervals between phrases, but did impact on the
pulse rate of chirping in section 2, pulse rate in the rolls, and rate of roll production
(P<0.05). The variation in calls among individuals for all variables analyzed was
2-7 times higher than the variation in calls within individuals. The frequency spec
trum of calls of male G. nigrijrons consisted of frequency bands of energy concen
trated between DC to 2000 Hz, with dominant frequencies at ca. 250 and 390 Hz.
In courtship, male calling began with section 1 type calls followed by section 2 and
section 3 type calls which were repeated until copulation. Females called only when
males produced section 2 and 3 calls. Calls of female G. nigrijrons consisted of low
frequency (dominant frequencies at 170 and 250 Hz) clicking sounds. Average dura
tions of female pulses were 15.4 msec (S.E. = 0.8, NIl) and average intervals of
778.1 msec (S.E. = 106.3, N = 1l)(Fig.2).
DISCUSSION
Leafhoppers and planthoppers produce vibrational signals by means of a sound
producing organ similar to the cicada timbal organs in the first and second abdomi
nal segments (Ossiannilsson 1949, Mitomi et at. 1984). The signals are transmitted
through the plant substrate on which the insect is sitting (Ichikawa and Ishii 1974,
Michelsen et al 1982). Males and females alternate calls and the male walks to the
female whereby copulation may occur. The mechanism for male mate location has
been described as phonoklinotaxis for the leafhopper, Amrasca devastans (Dis
t.)(Saxena and Kumar 1984). In contrast, Michelsen et al. (1982) theorized that
signal frequency-time domain information might be used to determine the direction,
distance, or both to a female. Claridge (1985) proposed that no directional informa
tion is received by the male, rather female calling elicits spontaneous random search
ing movements by males. Hunt (1988) proposed that male Graminella nigrijrons
receives information on distance but not direction to females. Additionally, Hunt
(1988) has demonstrated that phototactic responses in addition to acoustic signaling
facilitate male location of females.
When a male G. nigrijrons flies or hops to a plant it will produce a call spontane
ously. This signal, designated as a calling signal, attraction call, or common call
(Ossiannilsson 1949, Heady et al. 1986), is complex and consists of three distinct
sections (Fig. I). The G. nigrijrons call becomes more patterned over time. The first
portion of the call contains irregularly spaced clicks that increase in amplitude as
section 2 calls are produced. In section 3 the signal pattern is well defined. A similar
pattern of irregular clicking that changes to a patterned signal is found in the
leafhoppers Aphrodes trijasciatus (Fourer.) (Ossiannilsson 1949), Nephotettix
nigropictus (Dist.) (Claridge 1988), and short-winged forms of Macrosteles fasci
frons (Stal) (Purcell and Loher 1976). In the latter species, the male call was
described as a trill with gradual transition to chirping. However, other leafhoppers
12
..
VoL 24, No.1
2
I
I
2
3
UEC
c
0.5 SEC
Figure 1. Graminella nigrifrons male calling signal. (A) Entire calling signal composed of
three labeled sections. (B) Expanded Section 2 type calls with three repeated phrases. A phrase
(a) consists of chirps (b) and a roll (c). (C) Expanded Section 3 type calls with three repeated
phrases (a) consisting of chirps (b) and a roll (c). There are 6 pulses in the labeled roll.
studied do not produce a long building call, but rather, discrete chirps. Amrasca
devastans (Distant) was described as producing "croaking" sounds (Saxena and
Kumar 1984). Males of 10 Dalbulus species spontaneously produced calls composed
of chirps (Heady et al. 1986). For example, D. maidis (DeLong and Wolcott) emit
ted 1-11 repeated chirps spontaneously. Male common calls of 10 Dalbulus species
were analyzed using cluster analyses and the resulting phenogram was similar to
groupings of species based on cladistic analysis of morphological characteristics
(Heady et al. 1986). Although calls of members within a genus appeared similar in
Dalbulus, calls of members among related tribes do not seem similar using the
characteristics examined here. Dalbulus and Macrosteles, members of the tribe
Macrostelini, have very different calls. The call of G. nigrifrons is similar to N.
nigropictus, yet they are members of different Deitocephalinae tribes, Deltocepha
lini and Euscelini, respectively. Thus calls may be useful in resolving phylogenetic
relationships within genera but not at higher taxonomic levels.
Female calls of G. nigrifrons were very simple in structure compared to male calls,
and were produced only when males were producing section 2 and 3 signals. Section
1 male clicks do not elicit female calling nor any other obvious behaviors such as
walking, perhaps because they are not patterned and are not species specific, or
because their low amplitude may not be received by the female. Female calls of other
leafhoppers are also less complex (lacking different patterned sections) compared to
male calls. Examples are Dalbulus species (Heady et al. 1986; Heady, unpublished
data); Nephotettix species (Claridge 1988); M. jascifrons, whose female call was
described as a buzz (Purcell and Loher 1976); and A. devastans, whose call was
described as cooing (Saxena and Kumar 1984).
1991
13
••~ "; .i~'.Ii~.R"...~.n"'1
./'
./'
./'
./'
/'
\
1 SEC
\
\
\
/'
./'
\
./'
\
0.1 SEC
Figure 2. Graminella nigrifrons female calling signal produced during male section 2 or
section 3 signalling.
The differences in call complexity between males and females may have evolved as
a result of (1) the energetic cost of calling, (2) the risks associated with searching for
mates, and/or (3) runaway sexual selection. Integral to these three hypotheses is the
inherent asymmetry of reproductive roles between the sexes (West-Eberhard 1984).
Male leafhoppers, like many insects, produce large numbers of small gametes
(sperm) and multiply mate, whereas females invest more in gametes (large eggs),
produce relatively few of them, and mate once or twice. In most crickets, katydids,
and cicadas, males alone bear this energetically expensive activity of calling while a
mute female finds him (Alexander 1967). In leafhoppers, males bear most of calling
expense by producing more or longer calls than females.
Acoustic communication is the most energetically expensive of all forms of com
munication, i.e., chemical, visual, and tactile (Alcock 1989) and is known to be
exploited by predators and parasites due to the ease of locating the signaller. A
tachinid fly, Euphasiopteryx ochracea (Bigot), was attracted to taped songs of the
field cricket Gryl/us integer Scudder (Cade 1975) and a vertebrate predator (domes
ticated cat) was observed locating male crickets using their acoustic signals (Walker
1964). Additionally, the sarcophagid fly, Co/condamyia auditrix Shewell, acousti
cally intercepts calling male cicadas (Soper et al. 1976). Specific predators of the
leafhopper, G. nigrifrons have not been identified. Parasites identified from G.
nigrifrons include three pipunculid flies, one strepsipteran, two drynid wasps and
one encyrtid wasp (Freytag 1987); however, their host-finding strategies have not
been elucidated. Generalist insect and spider predators may be attracted to calling
male leafhoppers by detecting the vibrations on the plant and by cuing on the
14
Vol. 24, No. 1
movement of males as they walk to females. Predators and parasites probably exert
selective pressures on general male reproductive behaviors including song character
istics (Cade 1975, Zuk 1987a, b). Several studies of insect mating systems have
shown that males suffer greater predation and parasitism as a result of their conspic
uous displays and movements through habitats in search for females (Gwynne 1987,
Thornhill 1978, Burk 1982, Walker and Masaki 1989). Hunt (1988) found that male
G. nigrifrons flew plant-to-plant, calling on each, until landing on a plant that
harbored sedentary but calling virgin females. This "call-and-fly" tactic was previ
ously found in tick-tock cicadas, Cicadetta quadricincta and explained the higher
incidence of males than females caught in spider webs (Gwynne 1987).
Lastly, the complex calls of males could be the result of rapid or "runaway"
evolutionary change (Fisher 1958). If superior signalling is at a premium then there
is selection on females to favor superior signallers as mates, because they will
produce sons who are superior signallers. This leads to increasing selection on
signalling ability and a genetic correlation between female preference and male
signalling ability which will accelerate the evolution of both. Runaway change would
end when natural selection against too costly or too risky displays balances sexual
selection in favor of traits that are appealing to females (Alcock 1989). We expect
that male calls of G. nigrifrons contain different section types and are elaborate
compared to female calls due to female preference and yet are likely constrained by
the energetics of calling and the risks associated with mate-finding.
ACKNOWLEDGMENTS
We thank Peggy Rhee and Andrea Sweazy (College of Wooster), Rex Alvey
(ARS-USDA, OARDC-OSU), L.V. Madden (Dept. of Plant Pathology, OARDC
OSU), and The Borror Bioacoustic Laboratory, OSU for assistance. Salaries and
research support provided by State and Federal funds appropriated to the Ohio
Agric. Res. and Dev. Center, The Ohio State Univ. This is Journal Article No.
100-90.
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prey's calling song. Florida Entomologist 47:163-5.
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W. Loher (eds.), Cricket Behavior and Neurobiology. Cornell University Press.
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1991
17
NO INTERSEXUAL DIFFERENCES IN HOST SIZE AND
SPECIES USAGE IN SPALANGIA ENDIUS
(HYMENOPTERA: PTEROMALIDAE)
B. H. Kingl
ABSTRACT
Spa/angia endius were collected from fly pupae, primarily house fly and stable
fly, from a poultry house in Indiana. Male and female wasps did not differ within
and across host species in host size usage. Also, despite stable fly pupae being
significantly smaller than house fly pupae, the proportion of male wasps emerging
from the two host species was similar.
Early in the 1900's, entomologists observed that in some species of parasitoid
wasps, males tended to emerge from smaller hosts than did females, resulting in a
negative relationship between host size and parasitoid sex ratio (proportion males)
(reviewed in Flanders 1939, 1946). Since then, a group of sex ratio models, the host
quality models have been developed to explain this pattern (Charnov 1979, Charnov
et al. 1981, Werren 1984). These models were designed for solitary species (species in
which one offspring completes development per host). In these models, the predic
tion that male parasitoids should emerge from smaller hosts than females is based on
the assumption that developing on a small host will be more detrimental to a female
than to a male in terms of future ability to reproduce. The rationale of the assump
tion was that wasps will be smaller when developing on smaller hosts; and even small
males may be able to mate successfully, whereas small females probably lay fewer
eggs than large females (Charnov et al. 1981). There is some support for this idea in
a few species of parasitoid wasps (Charnov et al. 1981, Jones 1982, van den Assem et
al. 1989), but not in all species (King 1988). In those species or populations in which
the assumption is valid, natural selection is expected to favor females that oviposit a
greater proportion of males in small than in large hosts (Charnov et al. 1981).
Female wasps can potentially control the sex of their offspring by controlling fertil
ization because they have haplodiploid sex determination. Under haplodiploid sex
determination, males develop from unfertilized eggs and females from fertilized
eggs.
Here the host quality models' prediction of a greater proportion of sons from
small than from large hosts is tested using field collections of the solitary parasitoid
wasp Spa/angia endius Walker (Hymenoptera: Pteromalidae). The wasps emerged
from house fly pupae (Musca domestlea Linnaeus) and stable fly pupae (Stomoxys
ea/eitrans [LinnaeusJ) (Diptera: Muscidae) collected from a poultry house in north
ern Indiana.
IDepartment of Biological Sciences, Purdue University, West Lafayette, IN 47907.
Present address: Department of Biological Sciences, Northern Illinois University, DeKalb, IL
60115.
THE OREAT LAKES ENTOMOLOGIST
18
Vol. 24, No.1
Table I. -X + SD (n) size (mm 3) of house fly pupae from which male and female Spaiangia endius
emerged for five collection dates.
Fly Pupae Size
Date
Sep 03
Sep 10
Oct 01
Oct 08
Oct 15
Male wasps
26.24 ±
28.15
16.57 ±
19.31 ±
27.02 ±
3.09 (38)
2.27 (3)
5.04 (14)
7.19 (5)
3.21 (2)
Female wasps
25.86 ±
25.16 ±
18.51 ±
23.94 ±
25.81 ±
Test Comparison"
"'---:----~.
3.02 (110)
2.94 (14)
5.16 (30)
6.27 (14)
4.72 (15)
t = 0.66, P
t = 1.65, P
t = 1.17, P
t
1.37, P
0.35, P
=
=
0.51
0.12
0.12
0.09
0.73
*t-tests of whether females emerged from larger hosts than did males; two-tailed P values given when
means were in direction opposite to that predicted
Weekly from 28 May to 12 November 1985, fly pupae were collected from an
enclosed, shallow-pit, egg-layer poultry house in Delphi, Indiana (the Hilltop poul
try house described in Merchant 1984, Merchant et al. 1987), using pupal traps
(Hogsette and Butler 1981, Merchant et al. 1985). The pupae were held until flies
emerged and died. Then pupae from which no flies emerged and which had not been
depredated (Merchant 1984) were held individually in gelatin capsules for parasitoid
emergence. Wasp species were identified following Boucek (1963) and Rueda and
Axtell (1985) and house fly and stable fly pupae following Skidmore (1985).
Voucher specimens have been deposited at the museum of the Department of Ento
mology at Purdue University, West Lafayette, Indiana.
Width of fly pupae was measured to the nearest 0.05 mm at a magnification of
360 X under a dissecting microscope. These widths were converted to volumes (mm3)
using the following formulas: for house flies volume = (22.68) (width)-37.63 and
for stable flies volume
(11.31) (width)-11.18. These regression equations had
been determined by measuring both width and length on 17 stable flies and 36 house
flies, then calculating volume with the equation for a prolate spheroid (Holdaway
and Smith 1932), and finally regressing volume against width.
RESULTS
There were no significant differences among collection dates in S. endius sex ratio
> 0.50). Pooling across all dates, S. endius sex ratio was
26.1070 males (n = 330 male and female wasps).
There was no significant difference in the size of hosts from which male and
female S. endius emerged. This was true regardless of whether one looked (I) within
host species, (2) at all host species combined, or (3) between different host species.
Looking at house fly hosts. among the five dates with sample sizes of greater than
ten, a two-way analysis of variance on host size showed no significant effect of wasp
sex (F = 0.34, P = 0.56) and no significant interaction between wasp sex and date
(F = 2.26, P = 0.06). Because the interaction approached significance, I also did
individual t-tests for each date, but these also revealed no significant difference in
the size of hosts from which male and female wasps emerged (Table I). Looking at
stable fly hosts, combining all dates, there was no significant differen£.e in host size
between male and female wasps, though sample sizes are small (males: X ± SD 14.54
± 0.73, n = 4; females 14.50 ± 0.97, n 7; t 0.07, P = 0.94). Combining host
species, among the five dates with sample sizes of greater than ten, a two-way
analysis of variance on host size showed no significant effect of wasp sex (F = 0.51,
P = 0.48) and no significant interaction between sex and date (F = 1.57, P 0.18).
Looking between host species, though stable fly pupae are on average smaller than
(0 = 3.82, df = 6, P
1991
19
house fly pupae (Skidmore 1985, King 1991), the sex ratio of S. end/us on stable flies
(36010 males, n = 11 wasps) was not significantly greater than on house flies (25%
males, n
264 wasps) (G
0.67, P > 0.30).
DISCUSSION
The absence of host size differences between the sexes for S. endius in this study is
consistent with results of laboratory experiments by Donaldson and Walter (1984).
Their experiments indicated that S. endius females do not manipulate the sex of their
offspring in response to the size of house fly hosts. The lack of any significant
relationship between host size and parasitoid sex ratio for S. end/us contrasts both
with the pattern observed in most other species of parasitoid wasps that have been
examined and with the prediction of the host quality models. In most, though not
all, species of parasitoid wasps that have been examined, females emerge from larger
hosts than do males (about 44 of 65 parasitoid wasp species (reviewed in King 1987,
1989, in press).
Results with other species of Spalangia besides S. endius have been mixed. These
other species of Spaiangia are also parasitoids of fly pupae. Legner (1969) found the
predicted negative relationship between host size and offspring sex ratio for S.
nigra, but not for S. cameroni or S. drosophilae (statistical analyses of his results in
Table 7 of King 1987). King's (1988) laboratory results support the prediction for S.
cameroni parasitizing house flies. However, field results with S. cameroni are more
complicated: the prediction is supported on a within host species basis for two of
three dates - two dates on which S. cameroni emerged only from house fly pupae
but not on the date when stable fly pupae were also parasitized by S. cameroni (King
1991). Looking just at stable fly pupae, S. cameroni males actually emerged from
significantly larger hosts than did females, opposite the prediction. As was the case
with S. endius, S. cameroni sex ratio did not differ between small and large host
species (Le., stable fly and house fly pupae) (King 1991).
Because the members of the genus Spaiangia exhibit such a variety of relation
ships between sex ratio and host size, it is a promising group for a comparative
study. A comparison of behavioral, ecological, and life history factors among the
species may help to explain the interspecific differences in sex ratio patterns. One
possibility is that the species that do not support the host quality models' prediction
may not meet the models' assumption of a more positive effect of host size on
female than male reproduction. For example, competition among males for mates
may increase the importance of large male body size in some species. In general,
among parasitoid wasps, the importance of size on male reproductive success has
been little studied (Charnov et al. 1981, Jones 1982, King 1988, van den Assem et aI.
1989), especially relative to the attention given females (e.g., Charnovet aI. 1981,
Jones 1982, King 1988, van den Assem et al. 1989; 14 references in Table 6 of King
1987).
ACKNOWLEDGMENTS
I thank R. Flanders for introducing me to parasitoid wasps, the Brubakers of
Hilltop Egg Farm for the use of their poultry house, M. Merchant for helpful tips on
identification, R. Howard for valuable discussions in the planning, analyzing, and
writing of this research, and R. King for comments on the manuscript. This research
was aided by a National Science Foundation Graduate Fellowship and by Grants-in
Aid of Research from Sigma Xi, The Scientific Research Society.
20
Vol. 24, No. I
LITERATURE CITED
van den Assem, 1., J. J. A. van lersel, and R. L. Los-den Hartogh. 1989. Is being large more
important for female than for male parasitic wasps? Behavior 108:160-195.
Boucek, Z. 1963. A taxonomic study in Spalangia Latr. (Hymenoptera, Chalcidoidea). Acta
Entomo\ogica Musei Nationalis Pragae 35:429-497.
Charnov, E. L. 1979. The genetical evolution of patterns of sexuality: Darwinian fitness. Am.
Nat. 113:465-480.
Charnov, E. L., R. L. Los-den Hartogh, W. T. Jones, and J. van den Assem. 1981. Sex ratio
evolution in a variable environment. Nature 289:27-33.
Donaldson, J. S., and O. H. Walter. 1984. Sex ratios of Spalangia endius (Hymenoptera:
Pteromalidae), in relation to current theory. Eco!. Entomol. 9:395-402.
Flanders, S . E . 1939. Environmental control of sex in hymenopterous insects. Ann. Entomol.
Soc. Am. 32:11-26.
Flanders, S. E. 1946. Control of sex and sex-limited polymorphism in the Hymenoptera. Q.
Rev. BioI. 21:135-143.
Hogsette, J. A., and J. F. Butler. 1981. Field and laboratory devices for manipulation of pupal
parasites. pp. 90-94 In: Proceedings Workshop on Status of Biological Control of Filth
Flies. U.S.D.A.
Holdaway, F. T., and H. F. Smith. 1932. A relation between size of host puparia and sex ratio
of Alysia manducator. Aust. J. Exp. BioI. Med. Sci. 10: 247-259.
Jones, W. T. 1982. Sex ratio and host size in a parasitoid wasp. Behav. Eco!. Sociobiol.
10:207-210.
King, B. H. 1987. Offspring sex ratios in parasitoid wasps. Quart. Rev. BioI. 62:367-396.
___. 1988. Sex ratio manipulation in response to host size by the parasitoid wasp Spalangia
cameroni: A laboratory study. Evolution 42: 1190-1198.
___. 1989. Host-size-dependent sex ratios among parasitoid wasps: does host growth mat
ter? Oecologia 78:420-426.
___. 1991. A field study of host size effects on sex ratio of the parasitoid wasp Spalangia
cameroni. Am. MidI. Nat. 125:10-17.
Sex ratio manipulation by parasitoid wasps. In: Evolution and Diversity of Sex Ratio:
Insects and Mites (D. L. Wrensch and M. Ebbert, eds.). Chapman and Hall, New York, in
press.
Legner, E. F. 1969. Adult emergence interval and reproduction in parasitic Hymenoptera
influenced by host size and density. Ann. Entomo!' Soc. Am. 62:220-226.
Merchant, M. E. 1984. House flies and their parasitoids in egg-layer poultry houses in Indiana:
ecology and sampling method comparisons. M.S. thesis, Purdue Univ., West Lafayette,
IN.
Merchant, M. E., R. V. Flanders, and R. E. Williams. 1985. Sampling method comparisons
for estimation of parasitism of Musca domestica (Diptera: Muscidae) pupae in accumulated
poultry manure. J. Econ. Entomol. 16:716-721.
Merchant, M. E., R. V. Flanders, and R. E. Williams. 1987. Seasonal abundance and parasit
ism of house fly (Diptera: Muscidae) pupae in enclosed, shallow-pit poultry houses in
Indiana. Environ. Entomol. 16:716-721.
Rueda, L. M., and R. C. Axtell. 1985. Guide to common species of pupal parasites (Hymenop
tera: Pteromalidae) of the house fly and other muscoid flies associated with poultry and
livestock manure. Technical Bulletin 278. North Carolina Agricultural Research Service,
North Carolina State University, Raleigh, North Carolina. 88 pp.
Skidmore, P. 1985. The biology of the Muscidae of the world. Junk Publishers, Dordrecht,
Netherlands. 550 pp.
Werren, J. H. 1984. A model for sex ratio selection in parasitic wasps, local mate competition
and host quality effects. Neth. J. Zool. 34:81-%.
1991
21
A CROBASIS SHOOT MOTH (LEPIDOPTERA: PYRALIDAE) INFESTATION-TREE HEIGHT LINK IN A YOUNG BLACK WALNUT PLANTATION George Rinkl, Barbara C. Weber2, D. Michael Baines3, and David T. Funk4
ABSTRACT
Acrobasis shoot moth infestations were evaluated in a young black walnut prog
eny test for 4 years, from ages 3 to 6. Infestation levels were greatest on the largest
trees in the fourth and fifth year after plantation establishment, and were declining
by the sixth year. Acrobasis infestation appears to be a problem primarily on young
trees less than 2.5 m in height. There was no evidence for genetic resistance to
Acrobasis infestation in black walnut.
Black walnut (Jug/ans nigra) is commonly grown for production of high-quality
veneer logs. Only trees with stems free of form defects, such as forks or crooks in the
first 3 to 5 m, qualify as veneer logs. Stem defects in black walnut often result from
loss of apical dominance due to damage or mortality of the terminal bud, often
associated with late spring frosts. More recently, case bearer moths (Acrobasis spp.)
have been shown to damage terminal shoots and buds of young black walnut trees
when larvae tunnel into them. Our study was designed as a survey to assess damage
to trees in a young plantation with a relatively serious Acrobasis infestation. Insect
identification techniques followed those of McKeague and Simmons (1979).
METHODS
Open-pollinated seeds from 54 walnut trees were used to produce 1-0 seedlings for
a progeny test. Parent trees were selected from within a 250-mile semi-circular radius
south of the Shawnee National Forest in southern Illinois. This included an area
bounded by southern Illinois, western Kentucky, western Tennessee, and southeast
Kansas. However, most of the seed collections were from western Kentucky and
western Tennessee. In most cases seed was collected from one tree per stand.
Although trees of better than average form were sought at the time of collection, no
rigid minimum selection criteria were applied. Seedling progeny resulting from these
seed collections were planted in 5-tree row plots at a spacing of 1.8 m within rows
and 3.7 m between rows, in a randomized complete block design with 10 blocks;
upon establishment the plantation contained 2,700 trees.
The progeny test was established in early spring 1973 on a Haymond silt loam on
the floodplain of Sexton Creek, on the Shawnee National Forest in Alexander
IUSDA Forest Service, North Central Forest Experiment Station, Carbondale, IL 62901. 2USDA Forest Service. Washington. DC 20090. 3USDA Forest Service, Allegheny National Forest, Marienville, PA 16239. 4USDA Forest Service, Northeastern Forest Experiment Station, Durham, NH 03824. 22
Vol. 24, No. I
County, Illinois. The site was an abandoned pasture on which 1.2 m strips were
machine sprayed with a simazine, dalapon, 2,4-0 mix before planting and spot
sprayed in spring 1974 and 1975. In addition the plantation was mowed annually.
In 1976, after three growing seasons, we observed that many of the walnut trees
were growing slowly, some were crooked or forked, and a few had died back and
sprouted from near the base. We suspected insect attack as a possible cause of the
poor form and began surveys for the following insects: Acrobasis juglandis
(LeBaron), the pecan leaf casebearer; A. demotella Grote, the walnut shoot moth
(Martinat and Wallner 1980); and Xylosandrus germanus (Blandf.), an ambrosia
beetle. The ambrosia beetle was later associated with canker and dieback but not
with crook or forking problems related to terminal buds (Weber and McPherson
1984).
During 1977-1979, we made two types of surveys, using the same sampling
scheme for both. The surveys were done on a pilot basis in 1976 and repeated
1977-1979. First, during the dormant season, we counted I mm-Iong hibernacula
containing overwintering larvae of A. juglandis; the hibernaculum is the overwinter
ing case for the first instar larva. First instar larvae develop from eggs hatched in
late summer; they spin hibernacula in preparation for overwinter hibernation. Pres
ence of hibernacula on a tree was considered an indicator of susceptibility of the tree
to Acrobasis infestation. Second, in the early summer, we counted infested growing
shoots as a measure of damage by A. demotella; A. juglandis is a defoliator but does
not tunnel into shoots (Martin at and Wilson 1979). One tree per family-row plot per
block was randomly chosen (using a random number generator) for infestation
evaluation each fall; the same tree was evaluated the following spring. All insect and
infested shoot count data were square root transformed to increase normality. Tree
height was measured to the nearest em during the dormant season.
Data were analyzed by analysis of variance and covariance and correlation tech
niques. Family heritabilities were calculated using the H2 = (1-IIF) formula of
Kung and Bey (1979) in which F is the variance ratio for families.
During our 4-year evaluation of this plantation, height growth averaged 0.36 m
annually. Mean total tree height was 1.5 m in 1976 and the plantation average was
2.9 m by the end of the 1980 growing season. Infestation as reflected by the number
of hibernacula per tree increased from 1.4 in 1976 to a high of 2.6 in 1978 and
declined to 2.0 hibernacula per tree in 1979. Demographic trends established by
Acrobasis hibernacula were mirrored by the number of infested twigs per tree, which
increased from 1.3 in 1976 to 2.3 in 1978 and then declined to 1.7 in 1979 (table 1).
However, during the 1976-1979 period, the number of trees with hibernacula
increased from 20 percent in 1976 to 33 percent in 1977,44 percent in 1978, and 57
percent in 1979. Similarly, the number of trees with infested twigs increased from 20
percent in 1976 to 30 percent in 1977, 58 percent in 1978, and 63 percent in 1979.
Analyses of variance or covariance failed to disclose any significant family
effects. Heritability of susceptibility to infestation by Acrobasis was less than 0.15,
confirming that genetic resistance to Acrobasis damage is extremely low or nonexist
ent. Similarly there was no correlation between either number of hibernacula or
number of infested twigs and latitude or longitude of family origin.
Correlation analysis disclosed a pattern of very high positive correlations between
the number of hibernacula per tree and the number of infested twigs (0.87 < r <
0.97). These high correlations were expected and suggest that the parameters mea
sure similar attributes of Acrobasis infestation and that future surveys probably
need to measure only one of these parameters. The high correlations also suggest
that overwintering survival rates were high and parasitism/predator levels were
low.
1991
23
Table 1. - Average height (m), height growth (m), number of hibernacula per tree (no.) and
number of infested twigs
Mean
Minimum
Maximum
Height
Height
Height
Height
Height
Variable
76
77
78
79
80
1.5
1.8
2.1
2.4
2.9
0.2
0.4
0.4
0.6
0.8
2.9
3.6
4.4
5.5
6.1
Ht.
Ht.
Ht.
Ht.
76-77
77-78
78-79
79-80
0.35
0.26
0.35
0.46
-0.63
-1.15
-0.90
-2.55
1.39
1.67
2.78
2.02
Hib.76
Hib.77
Hib.78
Hib.79
1.4
2.0
2.6
2.0
1.0
1.0
1.0
1.0
7.0
14.0
16.0
3.0
Infest.
Infest.
Infest.
Infest.
1.3
1.7
2.3
1.7
1.0
1.0
1.0
1.0
15.0
3.0
14.0
9.0
Gr.
Gr.
Gr.
Gr.
76
77
78
79
Table 2. - Correlations of number of hibernacula and infested shoots with total tree height
(m) in current and subsequent years (ns = not statistically significant; •• = significant at P <
0.05)
Measurement
year
1976
1977
1978
1979
Tree height in
current year
Tree height in
subsequent year
Number of hibernacu/a
0.15 ns
0.18 ns
0.22 ••
0.23 ••
0.40 **
0.41
0.08 ns
0.07 ns
.*
Tree height in
current year
Tree height in
subsequent year
Number of infested shoots
0.04 ns
0.07 ns
0.30 ••
0.34 ••
0.45 **
0.43 **
0.04 ns
0.05 ns
A more interesting pattern of correlations emerged between number of hiberna
cula, infestation, and total tree height (table 2) and height growth (table 3). In 1976,
the first year of Acrobasis evaluation, there was no statistical correlation (r < 0.2)
between either measure of insect attack and growth variables. In 1977, there were
low but significant correlations (r > 0.2) of hibernacula and infestation with total
tree height. By 1978, both measures of insect attack were significantly correlated
with total height and height growth. Furthermore, the magnitude of the correlation
between total height and number of hibernacula doubled. However, by 1979 neither
measure of Acrobasis infestation was correlated with total height or with height
growth.
Such correlation patterns could result from the following circumstances: in 1976,
when there was no correlation between infestation and tree size or growth, the
24
Vol. 24, No. 1
Table 3. -Correlations of number of hibernacula and infested shoots with tree growth (m) in
significant at P < 0.05)
current and subsequent years (ns
not statistically significant; **
Measurement
year
1976
1977
1978
1979
Growth in
current year
Growth in
subsequent year
Growth in
current year
Number of hibernacula
0.14 ns
0.10 ns
0.13 ns
0.18 **
0.23 **
0.08 ns
0.20 **
Growth in
subsequent year
Number of infested shoots
0.08 ns
0.18 u
0.23 **
0.19 **
0.25 **
0.05 ns
0.04 ns
plantation was only 3 years old and all trees were in approximately the same size
category. The initial infestation was small and the outbreak was distributed among
trees of the same size class. By 1977 the outbreak was spreading to the larger trees as
evidenced by the positive corelation with tree height; by 1978, positive correlations
between infestation and both total height and height growth were present. By 1979,
on the basis of average number of hibernacula per tree or mean number of infested
twigs per tree, the infestation seemed to be declining. Furthermore, the remaining
Acrobasis no longer preferred the larger trees but infested any size trees.
All statistically significant correlations between total height and insect attack were
positive, suggesting that Acrobasis preferentially attacks larger trees once an infesta
tion is established in a plantation (table 2). Furthermore, it appears that in 1977 and
1978 Acrobasis also attacked the faster growing trees (table 3). The implication is
that, initially, Acrobasis infests the largest trees, those with the largest crowns;
however, in subsequent years when most of the trees in the plantation are in larger
size classes, insect attacks are of a more random nature.
The lack of negative correlations between infestation in a given year and growth in
subsequent years was surprising. Acrobasis damage has been described as resem
bling frost damage where infested terminals die back. In a large infestation, negative
correlations with growth in the following year might be expected. However, in
infestations at this population level, negative correlations were not encountered.
Instead, positive correlations imply that somehow infestations had little effect on
growth and perhaps were even slightly stimulatory. Thus, it is likely that the overall
infestation levels were not high enough to affect average growth rates.
By 1979, when the trees averaged 2.4 m in total height (table 1), infestation
appeared to be subsiding. Subsequent surveys did not disclose significant infesta
tions. Thus, we hypothesize that Acrobasis is a problem on young trees less than 2.5
m in height; stem form m'ly be affected but growth rate is not. Alternatively, a
buildup of populations of natural predators of Acrobasis may have resulted in a
declining Acrobasis population; unfortunately, no effort was made to monitor these
predators.
ACKNOWLEDGMENTS
We thank David J. Polak, for assistance with statistical analyses. At the time of
the study Dave Polak was a computer programmer with NCFES, Carbondale, IL.
We also acknowledge the cooperation of the Shawnee National Forest on whose
land the plantation was established.
1991
25
LITERATURE CITED
Kung, F. H. and C. F. Bey. 1979. Heritability construction for provenance and family selec
tion. In: Proceedings, Thirteenth Lake States For. Tree Impr. Conf., Aug. 17-18, 1977,
Univ. Minn., St. Paul, MN. USDA For. Servo Gen. Tech. Rep. NC-50, p. 136-146.
Martinat, P. J. and L. F. Wilson. 1979. The life histories of Acrobasis juglandis and A.
demotella on black walnut in Michigan. In: Walnut Insects and Diseases Workshop Proc.,
Carbondale, IL. USDA For. Servo Gen. Tech. Rep. NC-52, p. 40-43.
Martinat, P. J. and W. E. Wallner. 1980. Notes on the biology and damage of two Acrobasis
species (Lepidoptera: Pyralidae) on black walnut in Michigan. Great Lakes Entomol.
13:41-48.
McKeague, M. A. and G. A. Simmons. 1979. Terminal bud damage on black walnut. Michi
gan State Univ. Ext. Bull. E-1321. 3 p.
Weber, B. C. and J. E. McPherson. 1984. Attack on black walnut trees by the ambrosia beetle
Xylosandrus german us (Coleoptera: Scolytidae). Forest Science 30:864-870.
1991
27
HOST PLANT SUITABILITY AND A TEST OF THE FEEDING SPECIALIZATION HYPOTHESIS USING PAPILIO CRESPHONTES (LEPIDOPTERA: PAPILIONIDAE) J. Mark Scriber 1 and Robert V. Dowel12
ABSTRACT
The concept that host plant favorites would be used for more rapid and/or
efficient growth of the "locally adapted" individuals was tested in a preliminary way
using the giant swallowtail butterfly, Papilio cresphontes. Populations feeding only
on northern prickly ash, Zanthoxylum americanum (from Wisconsin), primarily (or
exclusively) on hoptree, Ptelea trijoliata (in Ohio) and on lime prickly ash, Z.
jagara, or Citrus, (in Florida) were compared on alternate hosts and on their actual
local hosts under controlled environmental conditions. While the results with final
instar larvae generally support the feeding specialization hypothesis with regard to
more rapid and/or more efficient growth on local food plant favorites, we are hesi
tant to extrapolate these results to the species as a whole for several reasons dis
cussed herein.
The giant swallowtail butterfly, Papilio cresphontes Cramer (Lepidoptera: Papi
lionidae) is a wide-ranging species, occurring from Minnesota and New England
south to the tip of Florida, westward to southern Arizona and California, through
out Mexico into Central America (Tyler 1975, Opler and Krizek 1984, Buetelspacher
and Howe 1984). The larvae, called "orangedogs", are occasional minor pests of
Citrus, but are also reported to feed on a large number of hosts from at least nine
plant families (Kimball 1965, Tietz 1972, Crocker and Simpson 1979). Many of these
literature records are suspect. However, it is certain that plant species in the Ruta
ceae are the favorite hosts for P. cresphontes and are the most frequently reported
(Scriber 1984). In fact, it has yet to be verified that the larvae of P. cresphontes can
grow and survive on any plants other than those in the Rutaceae family.
Within the Rutaceae there are local population preferences from northern prickly
ash Zanthoxylum americanum, in Wisconsin (Ebner 1970); hoptree, Ptelea trijoli
ata, in Illinois and Ohio and lime prickly ash, Zanthoxylum jagara, torchwood,
Amyris elemijera, and various Citrus spp. in Florida (Opier and Krizek 1984). The
Mexican Orange, Choisya dumosa, is a native Texas shrub which has also recently
been reported as a foodplant for P. cresphontes in Dallas, Texas (Crocker and
Simpson 1979).
The occurrence of local foodplant favoritism (Le. ecological monophagy) is com
mon in the tree-feeding species of Papilio in North America, and may in fact be the
rule rather than the exception in Lepidoptera. Scriber (1975) and Smiley (1978) have
suggested that such ecological monophagy might lead to increased biochemical and
lCurrent address and correspondence to: Dr. 1. Mark Scriber, Department of Entomol
ogy, Michigan State University, East Lansing, MI 48824.
2Current address: 1681 Pebblewood Dr., Sacramento, CA 95833.
28
Vol. 24, No.1
digestive efficiencies. Subsequent reduction of such efficiencies with respect to
ancestral or locally non-used (allopatric) foodplants might lead to what could be
called "obligate monophagy" (Smiley 1978). The evidence available for evaluation of
this concept is meager (see Fox and Morrow 1981, Scriber 1983, 1986).
The objective of this study was to assess the possibility that P. cresphontes popu
lations are differentially adapted to their local foodplant favorites compared to
other plants. Our approach was to collect P. cresphontes from various geographic
locations and carefully bioassay the larval growth performances on their local favor
ites as well as the allopatric foodplants which are local favorites used by other P.
cresphontes populations.
METHODS
Sources of Larvae
Larvae from eggs obtained from Citrus spp. in Broward County, Florida were
distributed at eclosion (unfed neonates) to the following foodplants: Lime prickly
ash, Zanthoxylum fagara; Northern prickly ash, Zanthoxylum americanum; sweet
orange Citrus sinensis Osbeck; and hoptree, Ptelea trijoliata. In Wisconsin, larvae
were obtained from eggs of a field-captured female in western Dane County and a
field-captured female in eastern Iowa County and subsequently were randomly
distributed to various rutaceous foodplants, including northern prickly ash and
hoptree. In these two populations (and nearly all of Wisconsin) prickly ash is the
only host plant available. For this study additional Wisconsin P. cresphontes were
obtained from Richland County where prickly ash, Z. americanum, shrubs are the
only hosts available. Ohio P. cresphontes were obtained from Preble County where
the local favorite and dominant host plant is hoptree, Ptelea trijoliata.
Feeding Experiments
Freshly molted penultimate instar larvae were individually weighed and distrib
uted to 150 x 25 mm petri dishes and maintained on a specific host under standard
ized rearing conditions (16:8 photo, scoto-phase with a corresponding 23:19-Co
thermoperiod, with moistened filter paper in each dish). Plant leaves collected from
several trees near the U. W. Arboretum or from our greenhouse were weighed fresh,
placed in water-filled floral aquapics® to maintain leaf turgor and subsequently
introduced into the petri dishes for standard gravimetric determination of food
consumption (Waldbauer 1968). Nutritional indices were calculated based upon the
dry weight (biomass) of leaves, feces, and larvae. The mean larval weight during the
stadium was estimated by the (initial plus final weight)l2. Indices of larvae perfor
mance are reported as in Scriber and Slansky (1981):
RGR, relative growth rate (mg biomass gained per day per mg larval biomass)
(RGR = RCR x AD X ECD)
RCR, relative consumption rate (mg biomass ingested per day per mg larval
biomass)
AD, approximate digestibility (also called assimilation efficiency) = Food infested (mg dry wt) - Feces (mg dry wt) 1000/1 Food ingested (mg dry wt)
x
0
BCD, efficiency of conversion of digested food (also called net growth effi
ciency) =
Biomass gained (mg dry 'wl) · x 1000/0
Food ingested (mg dry wt)-Feces (mg dry wt)
1991
29
ECI, efficiency of conversion of ingested food (also called gross growth effi
ciency) =
Biomass gained (mg dry wt) 1000/1
Food ingested (mg dry wt) x
0
ECI
AD x ECD = (overall efficiency)
Plants, larvae, and feces were frozen and freeze-dried for dry weight determina
tions. Statistical analyses were performed and where the ANOVA indicated signifi
cant differences between the means were analyzed by Tukey's test for unequal
sample sizes (Winer 1962, Snedecor and Cochran 1967). The very conservative
Tukey's test was used because our sample sizes were not as large as we had wished.
We therefore believe that all statistically significant differences represent very real
biological differences.
RESULTS
Growth of the Florida Papilio cresphontes, which uses only Citrus and lime
prickly ash as natural host plants, was significantly faster on lime prickly ash,
Zanthoxylum/agara, than was growth of Ohio P. cresphontes on this Florida plant
species for both the penultimate and the final instars (Table 1). The Florida popula
tion also had a higher efficiency of processing ingested biomass (ECI) than the Ohio
larvae. This central American plant species (Z. /agara) occurs only in the southern
half of Florida and southern tip of Texas (Fig. 1). As such this Ohio butterfly
population gets no closer than tOOO kilometers to the plant.
The congeneric northern prickly ash, Z. americanum, occurs sporadically from
Georgia and Alabama northward to the Great Lakes States and Canada. While all
Wisconsin populations and the Ohio populations are sympatric with northern
prickly ash and despite the fact that northern prickly ash is the only host plant for P.
cresphontes in Wisconsin, larval growth and efficiency of these larvae were no better
than for Florida larvae (which are totally allopatric with the plant range; Fig. 2 and
Table 2). No consistent differences are observed between the growth performances
of the penultimate and final instars of Wisconsin, Ohio, and Florida giant swallow
tail larvae. While Florida larvae exhibit the highest overall efficiency of processing
Z. americanum (ECI) in the penultimate instar, they exhibit the lowest efficiency in
the final instar (Table 2). The reverse is true for the consumption rates (RCR).
Ohio popUlations of P. cresphontes primarily use hoptree Ptelea trijoliata in the
source area (Preble Co., OH) for our experimental larvae (Fig. 3). Growth rates of
these Ohio larvae were nearly three times as fast (.215 mg mg- l dol) as Wisconsin
larvae in the final instar (Table 3). The Ohio efficiencies (ECI and ECD) averaged
twice those of Wisconsin larvae in the final instar (eg., ECI = 18.80/0 versus 8.1010
and 8.70/0 respectively). These patterns were not observed in the penultimate instar,
however the digestibility (AD) and consumption rates (RCR) were higher for Ohio
than Wisconsin larvae (Table 3). Not enough Florida larvae were available for
bioassays on hoptree.
DISCUSSION
Larvae of P. cresphontes from Florida, Ohio, and Wisconsin exhibit significant
differences in growth rates, consumption rates, and efficiencies of processing three
different plant species. For final instars, where the largest amounts of food are
consumed, populations in Florida grow faster and more efficiently on the local
foodpiant, lime prickly ash, than Ohio larvae (Fig. I). Similarly, populations in
Table I. -Growth performance of penultimate and final instar Papilio cresphonles fed Zanthoxylum jagara (lime prickly ash). Data are presented as a mean ±
SE. Instar
duration
(days)
(n)
Instar and Source
population
(5)
(4)
Ohio (Preble Co.)
Florida (Broward Co.)
Growth Rate
(RGR)
Consumption
Rate
(RCR)
6.6±0.2
6.1±0.1
n.s. .130±.01O
.224±.01O
16.2± 1.4
II.S±1.l
n.s.
.032±.003
.O84±.OO6
**
(A.D.)
(E.C.D.)
(E.C.I.)
Leaf quality
("10 water)
1.74±.13
2.16±.17
n.S.
24.3 ±4.4
43.0±4.S
33.9±4.5
26.0±4.9
n.s.
7.5 ±O.S
1O.5±0.9
69.2± 1.4
70.9±2.3
I.S9± .12
1.44± .41
n.s.
2S.S±2.9
27.2±3.1
n.s.
6.7±0.6
30.4± 14.3
n.s. 1.7±0.1
II.S± 1.1
66.7± 1.7
n.6± 1.6
•
w
c
~
Final Instar
Ohio (Preble Co.)
Florida (Broward Co.)
(3)
(3)
ISignificant differences are indicated ("
P±O.OS, ••
••
=
::cttl
0
:;c
ttl
P±O.OI) ANOVA (Snedecor and Cochran 1967).
~
t"'"
)
~
Table 2.-Growth performance of penultimate and final instar Papilio cresphontes fed Zanthoxylum americanum (northern prickly ash).
Instar and Source
population
Penultimate Instar WI (Dane Co.)
(Iowa Co.)
(Richland Co.)
OH (Preble Co.)
FL (Broward Co.)
(n)
(S)
(8)
(5)
(6)
(9)
L.S.D.I Final Inslar
WI (Dane Co.)
(Iowa Co.)
(Richland Co.)
OH (Preble Co.)
FL (Broward Co.)
L.S.D.I ISignifieant differences (P
(3)
(3)
(3)
(4)
(6)
Instar
duration
(days)
ttl
en
Growth Rate
(RGR)
Consumption
Rate
(RCR) (A.D.)
4.7±0.2 be
4.1±0.2ab
3.9±0.5 a
S.0±0.2 c
4.3±0.2 ab
.277 ± .011 bc
.321 ± .012 a
.307±.016 ab
.248±.017 c
.293 ± .005 abc
1.49±.06 a
1.35±.07 ab
1.34±.15 ab
l.30±.11 ab
1.l4± .05 b
36.0±2.2
3S.8± 2.1
46.2±3.3
42.3±S.4
39.7± J.3
(0.7)
(.049)
(.34)
(n.s.)
S.Sit 1.0 ab
8.1±0.S ab
a.s±o.s ab
7.0±0.0 b
1O.5±0.6 a
.I2S± .042
.135± .012
.107± .020
.160±.OOS
.100±.009
1.06± .28
1.04± .08
1.18±.17
1.27± .OS
1.67±.26
(2.8)
(n.s.)
(n.s.)
34.1 ±S.S
29.4±3.3
45.4±2.6
41.7±0.9
41.4±2.7
(14.3)
Efficiencies
(E.C.D.)
ab
b
a
ab
ab
Leaf
(E.C,!.)
(% water)
S3.6±4.2
63.0±3.8
56.2± 5.3
S1.8±9.9
66.3±3.6
IS.7±0.6 e
24.0±3.S ab
23.9±2.5 ab
19.3±0.9 c
26.0±0.8 a
71.1 ±0.9
70.8±0.7
n.S±0.9
74.0±0.S
77.8±0.1
(n.s.)
(4.4)
34.2± 2.4 b
45.4±S.1 a
20.0±0.8 c
30.4±1.7 b
IS.2± 1.2 c
11.4± 1.2 ab
13.0±0.4 a
9.0±0.4 be
12.6±0.4 a
6.3±0.7 c
(10.4)
(3.4)
ttl
Z
d
::0 t"'"
0
0
en
~
66.8± 1.4
69.9± 1.2
63.9± 1.4
73.2±0.9
74.4±4.1
O.OS) between the means are indicated by letters (Tukey's test for unequal sample sizes; Winer 1962, Snedecor and Cochran 1967).
<:
e.
~
Z
?
1991
31
7.5 %:t 2.8
Figure 1. Performance of final ins tar P. cresphontes larvae (data are presented as a mean ±
se) as a function of geographic source and range (shaded area) of the test plant, lime prickly
ash (Zanthoxylum Jagara).
Ohio grow faster and more efficiently on their local favorite, hoptree compared to
Wisconsin populations that prefer northern prickly ash (Fig. 3). Larvae of both
Ohio and Wisconsin populations grow faster and more efficiently than Florida
larvae on the northern prickly ash Z. americanum (Fig. 2).
While these results with final instars lend support to the feeding specialization
hypothesis and suggest that biochemical adaptation has occurred with local speciali
zation, the same patterns are not observed in all cases with larvae in their penulti
mate instar. While Florida larvae grow significantly faster and more efficiently on
Z./agara in the penultimate instar (Table 1), Ohio larvae in their penultimate instar
grow no faster on their favored Ptelea trifoliata (Table 2), and Wisconsin popula
32
Vol. 24, No.1
6.3 %:1:0.7
n=6
1\1111.£5
1,0 , 210 310!. 4,0
!
'0"
i
Figure 2. Performance of final instar P. cresphontes on northern prickly ash, Z. americanum
as a function of insect population and plant range (shaded).
tions do not grow significantly faster nor more efficiently than Florida larvae on
their favorite host Z. americanum (Table 3).
Variable leaf nutritional quality can be very important in determining the larval
growth rates and efficiencies (Scriber and Slansky 1981). Different growth perfor
mance on different plant species is not unexpected, but there are also significant
differences in nutritional quality within a plant species. For example, some leaf
quality variability (as indexed by leaf water content) may have been involved in the
differential growth performances to some extent in these studies, especially with
northern prickly ash where 8-100/0 differences in leaf water content were detected.
These differences may have been largely responsible for the good performance of
the Florida populations on northern prickly ash (Table 2).
Another important aspect of local adaptation (not addressed in this study) is the
critical first instar survival differences among geographic populations or host races
(Diehl and Bush 1984, Futuyma and Moreno 1988). Whether these differences in
survival may be related to behavioral (antixenosis) or toxicological (antibiosis)
mechanisms, they are of fundamental importance to the ecological success of the
1991
33
n=4
Hop-tree
f
tOO
Q
t!
•
I,!
MllES
200
I.!
300
I
401)
I
I!
,
Figure 3. Performance of final instar P. cresphontes larvae 0" noptree (Ptelea trifoliata) as a
function of the plant range (shaded).
population. The differences in growth performance detected at the penultimate and
final instars for surviving larvae may be of minimal significance compared to the
differential survival of the initial cohort of larvae in their neonate stages. However,
these later ins tars are around longer and other defenses appear to be useful (Fig. 4a,
4b).
For example, cryptic resting larvae on the side of the stern (Fig. 4a) could avoid
early bird detection and predation (Hirose et al. 1980), whereas osmeterial glands
(Fig. 4b) may be effective at deterring foraging ants or wasps as has been shown on
related Papilio in North America and Japan (Hirose and Tagaki 1980, Damman
1986). In addition, the resting spot for the larger larvae is on the inside of the thorny
w
"'"
-l
:t
Table 3. -Growth performance of penultimate and final instar Papilio cresphontes fed Pte/ea trifoliata (hoptree).
(n)
Instar and Source
population
t'I1
Instar
duration
(days)
Growth Rate
(RGR)
Consumption
Rate
(RCR)
(A.D.)
4.6±0.2 b
4.3±0.1 ab
3.8±0.2 a
.282±.020
.303 ± .009
.312± .030
1.29±.04 ab
1.21 ±.04 b
1.46± .05 a
42.8 ± 1.0 b
47.4±2.0 b
65.6±3.9 a
(0.7)
(n.s.)
(.180)
(7.6)
Efficiencies
(E.C.D.)
(E.C.I.)
Leaf
("10 water)
51.7±4.3 a
54.0±3.2 a
32.8±2.7 b
21.9 ± 1.6
25.3 ± 1.0
21.3 ± 1.3
71.6±0.4
71.6±0.4
n.9±0.4
(16.8)
(n.s.)
Penultimate Instar
WI
(Dane Co.)
(Iowa Co.)
OH (Preble Co.)
(9)
(8)
(3)
L.S.D.l
o
'"
E
r
>
~
t'I1
CIl
t'I1
Z
Final Instar
(3)
(5)
(4)
WI
(Dane Co.)
(Iowa Co.)
OH (Preble Co.)
L.S.D.l
I Significant
12.3 ± 1.0 b
9.9± 1.6 ab
6.3±0.1 a
.057±.007 b
.075±.012 b
.215±.005 a
0.71±.ll b
0.87± .08 ab
1.14±.0Ia
55.2±4.0
52.8±3.1
49.6±3.1
14.9±1.3 b
17.1 ±3.0 b
38.4±2.9 a
8.1±0.3b
8.7± l.l b
18.8±0.3 a
(5.20)
(.040)
(.31)
(n.s.)
(11.5)
(3.3)
69.3± 1.2
68.3 ± l.l
72.3 ±0.5
differences between the means (P = 0.05) are indicated by different letters (Tukey's test for unequal sample sizes: Snedecor and Cochran 1967, Winer
1962). (n.s. = nonsignificant differences via F-test, ANOVA)
d
3:
o
o
o
r
Ui
-l
<:
2..
.~
Z
o
'f
•
1991
35
Figure 4a-b. a-Final instar P. cresphontes resting cryptically on a stem of northern prickly ash,
Zanthoxylum americanum. b-Final instar "snake-like" osmeterial defensive response of P.
cresphontes larva to slight agitation. This has been shown for other Papilio to be a defensive
behavior to ants, wasps, birds or other predators (see text).
thick stems (Figs. 4a, 4b) down and away from the danger of browsing cattle (and
possibly deer) which were believed responsible for the pruned outer foliage on many
of these hoptree bushes and lower branches in Wisconsin. The use of taller trees of
prickly ash would avoid the mammalian grazing damage (Le., larvae consumed with
leaves). However, it has been shown that taller trees of Zanthoxylum have nutrition
ally poorer leaves upon which the Papilio growth is slower and mortality is higher
(Watanabe 1982). Female P. xuthus L. butterflies in Japan prefer these shorter
Zanthoxylum trees for oviposition (Watanabe 1982), as the P. cresphontes in Wis
consin also seem to do. Eggs and larvae of P. xuthus on Zanthoxylum in Japan in
early stages were attacked primarily by smaller predators such as ants, spiders, bugs,
and orthopteroids in contrast to birds and Polistes wasps that attack larger larvae
(Watanabe 1981).
Additional studies would be needed to determine whether the differences in larval
growth performance among the different P. cresphontes populations have a genetic
basis. The phytochemical cues for oviposition and the allelochemical factors making
the Rutaceae the primary host family of this insect are certainly of tremendous
ecological (Dethier 1941, 1954) and evolutionary significance for the Papilionidae
(Hancock 1983, Miller 1987). The specific chemical cues used by P. cresphontes for
oviposition and enhancement of larval feeding may not be unlike those for Japanese
Rutaceae feeders (Nishida 1977, Ichinose and Honda 1978). However, chemical
similarities in Rutaceae hosts with the Umbelliferae, Lauraceae, Aristolochiaceae,
Magnoliaceae, and Asteraceae (see Feeny et al. 1983) make the role of particular
allelochemics(and their interactions) in host selection and the ecology of different
Papilio uncertain at best.
36
Vol. 24, No. 1
This study reflects significant biological differences in larval performance among
different individual larvae on the same host plants under controlled conditions.
However we still lack the extensive geographic replication and intensive (intrapopu
lation) analysis to detect the extent of genetically based adaptations. We are there
fore hesitant to extrapolate the results from any particular host plant or any particu
lar population to the P. cresphontes species on the whole. Similarly, we have
illustrated that the relative differences in the roles of growth are in some cases instar
dependent. A more rigorous test of the feeding specialization hypothesis must
include not only controls for host plant nutritional quality, but should also include a
representative sample of genotypes from the entire range of the species of interest.
Our studies with the sympatric Papilio glaucus group are more extensive and inten
sive to date, and we have detected several different levels of behavioral biochemical
and genetic differences in the adaptation of local populations to favored host plants
(Scriber 1986, Scriber et al. 1989, 1990). Yet no single species can be used when
evaluating a general ecological concept such as the feeding specialization hypothesis.
A comparative approach using different taxa is essential, and thus additional data
from other insect/plant systems such as this study are still needed.
ACKNOWLEDGMENTS
This research was supported in part by the Michigan State University Experiment
Station (Projects MAES #1644 and 8072) and the College of Agricultural and Life
Sciences of the University of Wisconsin at Madison (Hatch #5134) and by a grant
from NSF (DEB 7921749). We thank Mark Evans and Nancy Teitelbaum for their
help in these studies.
LITERATURE CITED
Beutelspacher Baights, C.R. and W.H. Howe. 1984. Mariposas de Mexico. Fasc. l. La Prensa
Medica Mexicana, S.A. Mexico, D.F. 128 pp.
Crocker, R.L. and B.l. Simpson. 1979. Mexican orange, Choisya dumosa (Rutaceae), a poten
tial ornamental, is host for orangedog, Papilio cresphontes (Lepidoptera: Papilionidae).
Southwestern Entomol. 4:11-12.
Detheir, V.G. 1941. Chemical factors determining the choice of foodplants by Papi/io larvae.
Am. Nat. 75:61-73.
Dethier, V.G. 1954. Evolution of feeding preferences in phytophagous insects. Evolution
8:33-54.
Diehl, S.R., and G.L. Bush. 1984. An evolutionary and applied perspective of insect biotypes.
Ann. Rev. Entomol. 29:471-504.
Ebner, 1.A. 1970. Butterflies of Wisconsin. Milwaukee Public Museum Press. 205 pp.
Feeny, P., L. Rosenberry, and M. Carter. 1983. Chemical aspects of oviposition behavior in
butterflies. pp. 27-76 In: S. Ahmad, ed., Herbivorous Insects: Host-seeking behavior and
mechanisms. Academic Press, N.Y. 257 pp.
Fox, L.R. and R.A. Morrow. 1981. Specialization: Species property or local phenomenon?
Science 211 :877 -83.
Futuyma, 0.1. and G. Moreno. 1988. The evolution of ecological specialization. Ann. Rev.
Eco!. Syst. 19:207-233.
Hancock, D.L. 1983. Classification of the Papilionidae (Lepidoptera): a phylogenetic
approach. Smithersia 2:1-48.
Hirose, Y. and M. Tagaki. 1980. Attraction of two species of Polistes wasps to prey wounded
by them. Appl. EnL Zool. 15:108-110.
Hirose, Yo, Y. Suzuki, M. Tagaki, K. Hiehata, M. Yamasaki, H. Kimoto, M. Yamanaka, M.
Iga, and K. Yamaguchi. 1980. Population dynamics of the Citrus swallowtail, Papilio
1991
37
xuthus L. (Lepidoptera: Papilionidae): Mechanisms stabilizing its numbers. Res. Pop. Ecol.
21 :260-285.
Ichinose, T. and H. Honda. 1978. Ovipositional behavior of Papilio protenor demetrius Cra
mer and the factors involved in its host plants. App!. Entomo!. Zoo!. 13: 103-114.
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Miller, J.S. 1987. Host-plant relationships in the PapiJionidae (Lepidoptera): parallel clado
genesis or colonization? Cladistics 3:105-120.
Nishida, R. 1977. Oviposition stimulants of some Papilionid butterflies contained in their host
plants. Botyu-Kagaku 42:133-140.
Opler, P.A. and G.O. Krizek. 1984. Butterflies East of the Great Plains. Johns Hopkins Univ.
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Scriber, J.M. 1975. Comparative nutritional ecology of herbivorous insects: generalized and
specialized feeding strategies in the Papilionidae and Saturniidae (Lepidoptera). Ph.D. The
sis, Cornell University, Ithaca, New York. 283 pp.
___. 1983. The evolution of feeding specialization physiological efficiency, and host races.
pp. 373-412 In: R.F. Denno and M.S. McClure, eds., Variable Plants and Herbivores in
Natural and Managed Systems. Academic Press, N.Y.
1984. Larval foodplant utilization by the World Papilionidae (Lepidoptera): Latitudi
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___. 1986. Origins of the regional feeding abilities in the tiger swallowtail butterfly: ecologi
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71:94-103.
Scriber, J.M., R.L. Lindroth and J. Nitao. 1989. Differential toxicity of a phenolic glycoside
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Scribe.r, J .M., R. Hagen and R.C. Lederhouse. 1991. Foodplants and evolution within the
Papilio glaucus and Papilio troi/us species groups (Lepidoptera: Papilionidae). pp. 341-373
In: P.W. Price, T.M. Lewinsohn, G.W. Fernandes and W.W. Benson, eds., Plant-animal
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1991
39
GEOGRAPHIC DISTRIBUTION OF SIPHONAPTERA COLLECTED FROM SMALL MAMMALS ON LAKE MICHIGAN ISLANDS William C. Scharfl
ABSTRACT
The distribution of ten flea species collected from five small mammal host species
on 13 Lake Michigan islands is described. Four new eastern and southern records for
Hystrichopsylla dippiei Rothschild are given. Speculative suggestions are made
regarding dispersal routes of some of the small mammal host species, and the
distribution of flea species from Peromyscus maniculatus gracilis LeConte is dis
cussed in the context of island biogeography theory.
I collected fleas from small mammals on Lake Michigan Islands from 1%5 to
present. Two other previous studies from Lake Michigan islands (Hatt et. al 1948,
and Arnsman unpublished) each reported on flea species from one host species from
one island, and agree with my findings. I have verified the Arnsman specimens in
the Michigan State University Entomology Department collection. The Hatt et. al
specimens, residing in the University of Michigan Museum of Zoology collection
were identified by the eminent siphonapterist, Karl Jordan. A small portion of the
species listed here are included in Scharf and Stewart (1980), but most are identified
there by county and host only.
This listing is intended to shed light on possible distribution, and immigration
routes, to the islands by both the small mammals and their fleas beginning from the
melting of the Pleistocene ice sheets. I have previously speculated that the possible
mechanisms of small mammal distribution were by way of: (I) a land bridge during
the Lake Chippewa low water stage; (2) rafting on flotsam; (3) concealment in Indian
canoes or early settler's vessels (Scharf 1973, and Scharf and Jorae 1980). While
offering little conclusive evidence, the present study offers a few hints of the dispersal
pattern of both the hosts and fleas. This distributional evidence is then examined for
hypothetical extinctions of fleas due to island size and distance from the mainland in
the context of the island biogeography theory of MacArthur and Wilson (1967).
I collected 590 fleas of 10 species from 5 species of small mammal hosts on 13
islands in northeastern Lake Michigan (Figure 1). Capture was primarily by snap
trapping, but a live-trapping grid on High Island in June of 1986 and 1988
accounted for many fleas from that island. Firearms were used to collect squirrels
IDivision of Science and Mathematics, Northwestern Michigan College, Traverse City.
MI49684.
40
13
Vol. 24, No. 1
Squaw f..
~.
Ii:
Whiskey 1.•12~;!,a
Trout 1..:11
10,
Hig!J..I~
GullI. ~ty
.
f
en7
l,J?Hog I.
6
Beaver I.
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G
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L A
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I'"
&ale rimllee
Figure L Islands in Lake Michigan where small mammal fleas were collected. Numbers
correspond to Tables 1 and 2: 1- Marion Island; 2 - South Manitou Island; 3 North Manitou
Island; 4-South Fox Island; 5-North Fox Island; 6-Beaver Island; 7-Hog Island; 8
Garden Island; 9-High Island; iO-Guli Island; l l - Trout Island; 12- Whiskey Island; 13
Squaw Island.
during the hunting season. In most cases the mammals were brushed over a white
surface to find the fleas, which were preserved in 70070 ethyl alcohol, and mounted
on microscope slides in Canada balsam for identification. The flea specimens
remain in the private collection of the author. Skins and skulls of the hosts are in the
Northwestern Michigan College collection of vertebrates.
The major problem regarding collecting methods remains whether completeness of
seasonal, and habitat coverage is achieved, because some flea species show distinct
seasonal and habitat abundance. I sampled all of the islands with a trapping pressure
of at least 100 trap-nights. Some islands, such as the North and South Manitou
1991
41
Table I. - Flea species collected from Peromyscus maniculatus gracilis on Lake Michigan Islands.
Islands arranged from south to north, east to west and with numbers corresponding to Figure I.
FLEA SPECIES
Ctenophthalmus pseudagyrtl!!! Baker
Epetedia w. wenmanni (Rothschild)
Hyslrichopsylla dippiei Rothschild
Orchopeas leucopus (Baker)
Peromyscopsylla h. hesperomys (Baker)
X
X
2
3
4
5
ISLAND· 6 7 8 9
10
II
12
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
13 X
X
*I-Marion Island; 2-South Manitou Island; 3-North Manitou Island; 4-South Fox Island; 5
North Fox Island; 6-Beaver Island; 7-Hog Island; 8-Garden Island; 9-High Island; IO-GulI
Island; 11 - Trout Island; 12- Whiskey Island; 13 -Squaw Island.
Islands, South Fox Island, and High Island had over 2,000 trap-nights. These same
intensively trapped islands had the greatest seasonal span of trapping too, with early
spring to late fall trapping on the Manitou islands, and a wide span of summer to fall
on South Fox Island and High Island. Most of the specimens from Garden Island were
collected in August and September. The other islands were trapped between June and
August only, and the number of fleas collected per island reflect some degree of the
collecting effort. As a crude index of trapping pressure, I list the islands by increasing
number of fleas collected at each: Squaw, 8; Whiskey, 11; Gull, 12; Hog, 19; Marion,
28; High, 37; Garden, 40; Trout, 42; North Fox, 43; South Fox, 58; Beaver, 59; South
Manitou,79; and North Manitou, 154. Different habitat types, including hardwoods,
dunes and swamps were trapped on most islands.
Mammal names are according to Hamilton and Whitaker (1979). Most Siphonap
tera were identified using Holland (1949), but Benton (1983) was useful in a few
instances. The nomenclature of Holland is followed here.
The one mammal host species that is common to all of the islands visited is the
Woodland Deer Mouse, Peromyscus maniculatus gracilis LeConte, although
phenetic divergence has been shown between islands (Lederle et. al 1985). Table 1
shows the distribution of 5 flea species on this host by island and Figure 1 shows the
location of the islands keyed to the numbers in Table 1. Noteworthy for its ubiqui
tous distribution on this mouse is the flea Orchopeas leucopus (Baker) which is
found on every island.
The occurence of Histrichopsylla dippiei Rothschild from the deer mouse on four
islands (Table 1) is an important new range extension. This paper represents the
second Michigan record, and a new eastern and southern distribution for the spe
cies. While this flea has been noted on deer mice previously, the true host for this
species is considered by many to be the Red Squirrel, Tamiasciurus hudsonicus
(Erxleben), and the closest recorded occurrences of the flea are in Itasca and St.
Louis Counties in northeastern Minnestoa both on T. hudsonicus and P. manicula
tus (Timm 1975, and Benton 1980), and on P. maniculatus on Isle Royale (Nixon
and Johnson 1971). Its presence on the four Beaver islands closest to the Upper
Penninsula., and its distribution across Canada and the far northern U. S. provides
a clue to the route the host took to these islands. The absence of this flea from the
southern islands of the Lake Michigan archipelago indicates that small mammal
invasion after glacial retreat may have been by more than one route.
The other flea species collected from P. m. gracilis are commonly found on this
mouse, and widely distributed (Table 1). However, two fleas common on this host in
the U. P., Peromyscopsylla catatina (Jordan) and Megabothris quirini (Rothschild)
42
Vol. 24, No. 1
Table 2. - Flea species collected from Tamias striatus (L.) on Lake Michigan Islands. Islands
arranged from south to north. east to west and with numbers corresponding to Figure I.
FLEA SPECIES
2
Ctenophthalmus pseudagyrtes Baker
Megabothris acerb us (Jordan)
Tamiophila grandis (Rothschild)
x
3
ISLAND*
4
6
8
X
X
X
X
X
X
X
* 2-South Manitou Island; 3-North Manitou Island; 4-South Fox Island; 6- Beaver Island; 8
Garden Island;
are notably absent from the Lake Michigan island mice (Lawrence et. al 1965,
Scharf and Stewart 1980, Scharf et al. 1991).
The distribution of fleas from P. m. gracilis on the Lake Michigan Islands contra
dicts both the distance effect, which predicts that on islands of similar size the
number of species vary inversely with the distance from the mainland, and the size
effect which predicts that species numbers vary directly with the area avialable for
immigration as proposed by the island biogeography theory of MacArthur and
Wilson (1967). First, the non-conformity with the distance effect is illustrated in
Table I, by Marion Island, a relatively small island within 3 km of the mainland
having as few species of fleas from this host as Squaw and Whiskey Islands, which
are similar sized islands, but which are more than 15 km from the mainland. This is
further shown by Gull Island, which is the most isolated of the islands, and has one
more flea species than the three previous islands. Trout Island, of about the same
size and distance from the mainland, is tied for the greatest number of flea species
from this host for any of the 13 Lake Michigan islands studied.
Second, the size effect shows a lack of conformity with the theory because the
largest islands, North and South Manitou Islands, North and South Fox Islands,
and Beaver Island have only three and four species each. This is fewer species than
the 5 species of either Hog Island or Trout Island which are considerably smaller. In
addition, Garden Island with only 2 species, and one of the largest islands studied
has only 2 recorded species of fleas and is within 4 km of Trout and Hog Island
which have the most flea species.
I recognize several reasons for caution in using these data from the mouse flea
species and islands to measure the validity of the island biogeography theory of
MacArthur and Wilson (1967). These are: (1) I have no certainty that I have collected
all the flea species from each island; (2) I have no indication of what levels of
equilibrium between extinction and immigration are in these islands, much less
whether those levels have been reached or not; (3) the area effect, where an increase in
area lowers extinction, seems to be unrelated to the sizes of the islands in this study;
and (4) the distance effect, lowering immigration in direct proportion to the distance
from the mainland, varies depending on whether a person hypothesizes immigration
from the Wisconsin side, the Upper Peninsula or the Michigan Mainland.
The distribution of flea species from the Eastern Chipmunk Tamias striatus (L.) is
given in Table 2. Megabothris acerbus (Jordan) is found on all 5 islands where T.
striatus was collected. Tamiophila grandis (Rothschild) is found only on this host on
South Manitou Island and South Fox Island, and was recorded from the Fox Squir
rel Sciurus niger (L.) from North Manitou Island. The only other Michigan records
of this flea are from chipmunks on the mainland of Leelanau County (Scharf and
Stewart 1980). This indicates that the colonization route may have been from the
adjacent mainland for this host on the southern islands of the archipelago. Chip
munks are absent from most of the islands where we failed to collect them with the
exceptions of Beaver, Marion, and North Fox islands.
The other four hosts and their fleas are listed here as unremarkable occurrences
both because of the limited number of islands inhabited by each mammal species,
1991
43
and the ubiquitous distribution of their fleas. The Red-backed vole, Clethrionomys
gapperi (Vigors), harbored Ctenophthalmus pseudagyrtes Baker on North Manitou
Island, and Peromyscopsylla h. hesperomys (Baker) on Beaver Island. The Meadow
Vole, Microtus pennsylvanicus (Ord) was parasitized by C. pseudagyrtes on Marion
Island (Scharf 1984). The Fox Squirrel Sciurus niger (L.) had Orchopeas howardii
(Baker) on North Manitou Island in addition to the chipmunk flea noted above. I
also collected Corrodopsylla c. curvata (Rothschild) from the Masked Shrew, Sorex
cinereus Kerr, on North and South Manitou Islands.
ACKNOWLEDGMENTS
I thank the William R. Angell Foundation for financial support through their
grants to Northwestern Michigan College for island biology studies during over 26
years of Great Lakes island studies. Cooperation from the Michigan Department of
Natural Resources, Wildlife Division, on High and Garden Islands, the U. S. Fish
and Wildlife Service on Gull Island, and the National Park Service on the Manitou
Islands is gratefully acknowledged. The following persons in addition to numerous
generations of NMC Natural History of Vertebrates and Ecology students helped
trap and collect fleas: T. Allan, B. Busch, M. Fitch, 1. Haswell, G. Hansen, M.
Hindelang, P. Lederle, L. Osterlin, 1. Mason, 1. Rogers, G. Shugart, V. Shugart, E.
Scharf, K. Scharf, K. Stewart, W. Wagoner, and R. Zillman. I also thank Dr. Omer
R. Larson for early encouragement in the study of Siphonaptera.
LITERATURE CITED
Benton, A. H. 1980. An atlas of the fleas of the eastern United States. Marginal Media,
Fredonia, N. Y.
Benton, A. H. 1983. An illustrated key to the fleas of the eastern United States. Bioguide No.3
Marginal Media, Fredonia, N. Y.
Hamilton, W. J. and J. O. Whitaker. 1979. Mammals of the Eastern United States. 2nd ed.
Cornell Univ. Press, Ithaca, N.Y.
Hatt, R. T., J. Van Tyne, L. C. Stuart, C. H. Pope, and A. B. Grobman. 1948. Island Life: a
study of the land vertebrates of eastern Lake Michigan. Cranbrook Inst. Sci. Bull.
27:1-179.
Holland, G. P. 1949. The Siphonaptera of Canada. Canadian Dept Agric. Bull. 70.
Lawrence, W. H., K.L. Hays, and S. A. Graham. 1965. Arthropodous ectoparasites of some
northern Michigan mammals. Occas. Papers Mus. Zool. Univ. Michigan. 639:1-7.
Lederle, P. E., W. C. Scharf, G. W. Shugart, and M. Fitch. 1985. Size variation in Peromyscus
maniculatus gracilis from the Beaver Islands. Jack-pine Warbler 63:107-110.
MacArthur, R. H. and E. O. Wilson. 1967. The Theory of Island Biogeography. Princeton
University Press, Princeton, N.J. xi + 203 p.
Scharf, W. C. 1973. Birds and land vertebrates of South Manitou Island. Jack-pine Warbler
51;2-19.
Scharf, W. C., and M. L. Jorae 1980. Birds and Land Vertebrates of North Manitou Island.
Jack-pine Warbler 58:4-15.
Scharf, W. C. and K. R. Stewart. 1980. New records of Siphonaptera from northern Michigan.
Great Lakes Entomol. 13:165-167.
Scharf, W. C. 1985. Meadow voles on Marion Island. Jack-pine Warbler 62: 77.
Scharf, W. C., P. E. Lederle, and T. A. Allan. 1991. Siphonaptera from the central and eastern
Upper Peninsula of Michigan. Great Lakes Entomol. 23:201-203.
Timm, R. M. 1975. Distribution, natural history and parasites of mammals of Cook county,
Minnesota. Occas. Papers, Bell Mus. Nat. Hist. 14:1-56.
Wilson, N. and W. J. Johnson. 1971. Ectoparasites of Isle Royale, Michigan. The Michigan
Entomo!. 4:109-115.
1991
45
ACORNS AS BREEDING SITES FOR CHYMOMYZA AMOENA (LOEW) (DIPTERA: DROSOPHILIDAE) IN VIRGINIA AND MICHIGAN Henretta Trent Band I
ABSTRACT
Chymomyza amoena is the only chymomyzid fly emerging from white oak acorns
in Virginia. An average of 2-3 adult flies emerged from a single acorn in July while
emergence declined to 0.4 adults/acorns in September. In fall, Drosophila melano
gaster was also present. The incidence of drosophilid (Drosophila, Chymomyza)
larvae in parasitized acorns in Virginia (400/0) in autumn was significantly greater
than in Michigan (14%). The Chymomyza larvae present in the parasitized acorns in
Michigan most likely were C. amoena, from the known adaptation of this species in
Michigan to frass-breeding.
Williams (l989a) has compiled a list of North American nut-infesting insects and
their host plants. Chymomyza amoena (Loew) was among the insects contributing to
northern red oak (Quercus rubra) acorn decay in Illinois (Winston 1956) and larvae
were found in acorns of mixed oak Quercus species in West Virginia (Dorsey et al.
1962). Drosophilids do not everywhere use the same breeding substrates. For instance,
Drosophila pseudoobscura Frolowa and D. persimilis Dobzhansky and Epling breed
in slime fluxes at Mather, California (Carson 1951, Carson et al. 1956) but not at
Blodgett Forest 127 km distant (Spieth 1987). The rarity of slime fluxes led Spieth
(1987) to discover that drosophilids could breed in California black oak Quercus
kelloggii acorns, if moist. Drosophila subobscura Collin in England breeds in rowan
berries but not elderberries; the reverse occurs in Switzerland (Burla et al. 1987).
Chymomyza amoena breeds in a variety of fruits in Michigan, including domestic
(commercial) apples Malus pumilia, in apples in Virginia and in black walnuts
Juglans nigra in Michigan (Band 1988a,b,c,d, 1989a). Larvae have been found to
overwinter in a variety of substrates in Michigan (Band and Band 1984, 1987) and in
apples in Virginia (Band 1988a). The question arises; does this species also breed in
acorns in the two states? Here I demonstrate C. amoena emergence from acorns in
Virginia. Larval adaptation to frass breeding arising from female exploitation of
parasitized substrates for oviposition (Band 1988a,b,d, 1989a) suggests that C.
amoena is also the chymomyzid in acorns in Michigan.
Studies in Virginia were carried out at Mountain Lake Biological Station (MLBS),
elev. approx 1200 m, Giles Co. Virginia, during the summer and fall of 1989 and at
IDepartment of Zoology. Michigan State University, East Lansing, MI 48824.
46
Vol. 24, No. 1
two sites in Michigan in the fall of 1989. The 600 acre MLBS site with mixed
deciduous and evergreen forests is adjacent to the 100,000 acre Jefferson National
Forest. Other Chymomyza and Drosophila species occur in Giles Co., Virginia
(Band 1988c,d, 1989b). Chymomyza eggs characteristically have a crown of short
filaments (Sturtevant 1921, Throckmorton 1%2, Schumann 1987). Larvae can be
distinguished by the posterior spiracular region (Hackman et al. 1970). To date, C.
amoena is the only chymomyzid consistently recorded in frass or frassy interiors
(Dorsey et al. 1962, Band 1988a).
Study sites in Michigan were stands of oaks. In the northern lower peninsula the
site was between Gaylord and Grayling adjacent to US Interstate route 75 and in
mid-Michigan the site was a stand of oaks near a housing development in Meridian
Twp., Ingham Co. off Grand River Ave. Chymomyza procnemoides has also been
collected in Michigan as well as Virginia (Wheeler 1952) but its breeding sites are
unknown.
Virginia. White oak acorns identified as Quercus alba were collected at two
locations near the Station on 12 July and 25 July 1989. The acorns collected were
rejects among piles of consumed acorns abandoned by squirrels or were unpiled
acorns lying 1-2 m adjacent to a foot trail through the woods. All acorns were
microscopically inspected for the presence of Chymomyza larvae and/or eggs and
tentatively identified as C. amoena. Larvae were reared in a glass population jar
over moist paper toweling. Some of the emerging C. amoena adults were transferred
to apple + frass to determine if these acorn flies were comparable to C. amoena
emerging from other substrates (Band 1988a,b) and could utilize apples as a food
source; the rest were maintianed on high protein laboratory medium (Band 1988a).
On 30 September 1989 acorns were more rigorously collected on the ground in
order to assess the frequency of those damaged and those infested by drosophilid
larvae. All were transported back to Michigan State University where acorns with
holes were separated, dissected and inspected for the presence of drosophilid larvae
or eggs.
Michigan. On 16 October 1989 large numbers of red oak acorns were collected at
a site in northern lower (NL) Michigan in Crawford Co. and on 20 November in
mid-Michigan in Ingham Co. (EL). Both groups were sorted, and examined follow
ing the same protocols as those for the Virginia Fall season collection.
Data analysis. G-statistics have been calculated according to Sokal and Rohlf
(1969).
RESULTS
Virginia. Chymomyza amoena was the only chymomyzid to emerge from the
infested acorns. Eleven of 16 acorns collected in mid-July contained 31 C. amoena
larvae and 16 of 20 acorns collected in late July had 133 eggs and larvae. Eggs were
found in the vascular elements at the base of the acorn or inside on frassy pulp, as
found by Spieth (1987) for drosophilid oviposition in acorns in California. How
ever, no other drosophilids were detected in summer. Table 1 records the numbers of
adults emerging from the July and September MLBS acorn collections. Adults
emerging readily oviposited on apples + frass; 76 adults emerged and produced an
F2. Ten acorn emergents showed the ability to switch to a fruit substrate and retain
fertility, as expected (Band 1988a). As observed for C. amoena egg counts in apples
(Band, 1988a), egg totals in acorns exceeded the numbers of adults emerging.
Twenty-two percent of the acorns collected in September in Virginia were dam
aged (n = 216) of which 400/0 had drosophilid larvae inside. No acorns collected in
September contained eggs on the outer surface of damaged or undamaged acorns.
Only parasitized acorns were used for drosophilid oviposition in September among
the newly fallen acorns but the initial parasite, probably a curculionid larva, had
1991
47
Table I.-ChymomyZa amoena emergence from damaged white oak acorns in Virginia in July and
September 1989.
Month
No. infested
acorns
Emerged
species
C
C.
D.
C
11
16
17
No. adults
emerged
Ave. no. adults
emerged per acorn
24
50
4
2.2
3.1
0.2
0.4
amoena
amoena
melanogaster
amoena
7
Table 2. -Comparison of acorns infested with Chymomyza amoena and other drosophilid larvae
from Mt. Lake Biological Station (MLBS), VA, Northern Lower (NL) Michigan and East Lansing
(EL), MI in Fall, 1989.
Acorns
State
Virginia
Michigan
Michigan
Location
MLBS
NL
EL
Total
216
210
327
among damaged acorns:
No. Damaged
Undamaged (010)
(79010)
(95010)
(92010 )
168
200
303
2; G
No
larvae
With
larvae
(010)
Total
29
7
22
19
3
2
(40010)
(30010)
( 8010)
48
10
24
8.72; P 0.05
exited. Five Drosophila larvae and 28 C. amoena larvae were counted in the 19
acorns; 4 D. melanogaster and 7 C. amoena adults emerged.
Michigan. Six percent of the total red oak acorns collected (n
537) were
damaged with 14% of these containing drosophilid larvae (Table 2). These infesta
tion rates are significantly lower than that encountered in Virginia. The intervening
month between collecting in northern lower Michigan and in mid-Michigan did not
increase the rate of drosophilid acorn infestation in mid-Michigan despite area wide
C. amoena overwintering/substrate utilization studies in past years which estab
lished its presence in the greater Lansing/East Lansing area and adjacent communi
ties (Band and Band 1984).
Michigan C. amoena females also oviposited only in parasitized acorns. As in
Virginia in September, no 'eggs or remnants of egg cases were detected in vacular
elements at the base of the acorns. There were 1 Drosophila and 4 C. amoena larvae
in the 3 NL acorns, 5 C. amoena larvae in the 2 EL acorns.
DISCUSSION
Sturtevant (1921) was the first to report that C. amoena bred in a variety of fruits
and nuts: Banks bred it from acorns, Shannon from walnut and butternut husks, he
and Metz obtained it from apples and bananas (p. 61). Winston (1956) reared all
larvae to adulthood, thereby determining that C. amoena bred in parasitized and
decaying acorns. He labeled them fungal feeders and scavengers. Dorsey et al.
(1962) found that curculionid larvae were the primary acorn pests in West Virginia
forests and listed C. amoena among the secondary invaders. Sampling over a year's
time, beginning in September, Winston (1956) found that larval-infested acorns were
rejected by squirrels. Weckerly et al (1989) discovered that intact acorns from which
curculionid larvae had not emerged were consumed by squirrels. Winston (1956)
also found that moisture was important, as Spieth (1987) did later.
Since discovering that C. amoena in mid-Michigan was using such substrates as
unripe firm apples for larval development in summer (Band 1988a,b) and firm
48
Vol. 24, No.1
native crabapples Malus coronaria for overwintering (Band and Band 1984), my
interests have included comparative substrate use and how females are able to
oviposit in firm substrates. Numbers emerging from acorns versus initial numbers of
eggs and larvae show a sharp reduction as in apples (Band 1988a). Exit holes made
by initial pest larvae (Moffett 1989, Williams 1989b) provide an entrance way,
analogous to pest larvae breaking the surface in a developing apple. For Virginia,
these findings indicate that the surrounding forest may be the reservoir from which
C. amoena can invade fallen and un fallen apples yearly. Two major summer sites
which have larval-infested apples lack fallen apples by late Fall (Band 1988c). The
ability to invade and breed in acorns damaged by primary insect attackers indicates
an inter-species dependency that may contribute to the survival of this species in
forested and urban areas. The Virginia (reported here), West Virginia (Dorsey et al.
1962) and Illinois (Winston 1956) studies also indicate C. amoena was a forest!
woodland species where it evidently was breeding in such native nuts and fruits as
acorns, black walnuts, and endemic crabapples before the colonists arrived.
The significantly lower numbers of infested Michigan acorns as compared with
Virginia may reflect a lower population density and/or totally migrant female ovi
position. Neither site is immediately adjacent to areas in northern lower Michigan or
mid-Michigan used in past studies on overwintering/substrate utilization (Band and
Band 1982, 1984). Biotic factors as availibility of substrates and abiotic factors as
altitude, latitude and past glaciation history influence species composition. Oviposi
tion behavior on fruits in summer has been strikingly similar in Michigan and
Virginia (Band 1989a).
The presence of D. melanogaster in acorns in Virginia is not unexpected. Dro
sophila species can breed in parasitized acorns in California (Spieth 1987) and have
emerged from frass in Hawaii (Heed 1968). Three species have adapted to the
nephritic gills of crabs (Carson 1974).
To date however, C. amoena has been the only chymomyzid in parasitized acorns,
a female oviposition behavior and larval developmental adaptation that may have
enabled this species to follow other pests into unripe parasitized domestic fruits.
Breeding sites and breeding behavior of this species in Europe (Bachli and Rocha
Pite 1982, Schumann 1987) remain unknown. Breeding sites for other forest Chy
momyza in the East remain unknown.
NOTE ADDED
In a letter dated 16 Oct. 1990, Dr. Hans Burla, Zoological Museum, University of
Zurich, reported that he had bred 10 females and 14 males of C. amoena from 100
chestnuts, Casatanea sativa (= vulgaris = vesica), collected in a forest in the Ticino
Canton of Switzerland. The chestnuts were old, and had been collected in the forest,
not far from the edge. The shipment also contained crabapples, apples, pears and
plums. Thirteen C. amoena adults emerged from the 13 apples.
Dr. Burla verified that this species of Castanea is native to Europe. This again
revives the comment initially made by Bachli and Rocha Pite (1981) after C. amoena
was collected in eastern Europe, that this species may be introduced, and capable of
spreading under favorable conditions as C. procnemis did in Japan (Okada 1976) or
may indicate that C. amoena has a Holarctic ditribution.
ACKNOWLEDGMENTS
Appreciation is gratefully extended to Jim Murray, Director, University of Virgin
ia's Mountain Lake Biological Station for providing research space in summer, 1989.
I thank Spencer Tomb for identifying the Quercus alba acorns and Jon Beaman the
1991
49
Quercus rubra acorns. Reviewers' comments have been helpful in revising the
manuscript.
LITERATURE CITED
Biichli, G. and M.T. Rocha Pite. 1981. Drosophlidae of the Palearctic Region, pp. 169-196.
In: M. Ashburner, H.L. Carson and J.N. Thompson, Jr.(Eds.). Genetics and biology of
Drosophila. Vol. 3a. Academic Press, N.Y. 429 + Ixx pp.
Biichli, G. and M. T. Rocha Pite. 1982. Annotated bibliography of Palearctic species of
Drosophilidae (Diptera). Beitr. Entomol. 32:303-392.
Band, H. T. 1988a. Host shifts of Chymomyza amoena (Diptera: Drosophilidae). Amer. Mid.
Nat. 120:163-182.
___. 1988b. Chymomyza amoena (Diptera: Drosophilidae): an unusual urban drosophilid.
Va. J. Sci. 39:242-249.
_ _. 1988c. Chymomyza amoena (Diptera: Drosophilidae) in Virginia. Va. J. Sci.
39:378-392.
1988d. Behavior and taxonomy of a chymomyzid fly (Chymomyza amoena). Intern.
1. Compo Psycho!. 2:3-26.
_ _. 1989b. Aggregated oviposition by Chymomyza amoena (Diptera: Drosophilidae).
Experientia 45:893-895.
_ _. 1989b. Behavior of the Chymomyza aldrichii species group (Diptera: Drosophilidae)
in Virginia's Allegheny Mountains. Va. J. Sci. 40: 230-237.
Band, H. T. and R. N. Band. 1982. Multiple overwintering mechanisms by Chymomya
amoena (Diptera: Drosophilidae) in Michigan and laboratory induction of freeze tolerance.
Experientia 38:1448-1449.
1984. A mild winter delays supercooling point elevation in freeze tolerant Chymomyza
amoena larvae (Diptera: Drosophilidae). Experientia 40:889-891.
_ _. 1987. Amino acid and allozyme changes in overwintering Chymomyza amoena (Dip
tera: Drosophilidae) larvae. Experientia 43: 1027-1029.
Burla, H., H. Kiinzli, and U. Pfaendler. 1987. Laboratory experiments on natural breeding
substrates in Drosophila subobscura. Genet. Jber. 39:269-285.
Carson, H. L. 1951. Breeding sites of Drosophila pseudoobscura and D. persimilis in the
transition zone of the Sierra Nevada. Evolution 5:91-96.
1971. The ecology of Drosophila breeding sites. The Harold L. Lyon Arboretum
Lecture, University of Hawaii Foundation.
___. 1974. Three flies and three islands: Parallel evolution in Drosophila. Proc. Nat. Acad.
Sci. 71:3517-3521.
Carson, H. L., E. P. Knapp and H. J. Phafr. 1956. The yeast flora of the natural breeding sites
of some species of Drosophila. Ecology 37: 538-543.
Dorsey, C. K., E. H. Tryon, and K. L. Carvell. 1962. Insect damage in acorns in West Virginia
and control studies using granular systemic insecticides. J. Econ. Entomo!. 55:885-888.
Hackman, W., S. Lakovaara, A. Saura, M. Sorsa and K. Vepsalainen. 1970. On the biology
and karyology of Chymomzya costata Zetterstedt with reference to the taxonomy, and
distribution of various species of Chymomyza (Diptera: Drosophilidae). Ann. Entomol.
Fenn. 36:1-9.
Heed, W. B. 1968. Ecology of the Hawaiian Drosophilidae. Univ. of Texas Pub!. No.
6818:387-419.
Moffett, M. W. 1989. Life in a nutshell. Nat. Geographic Mag. 175:783-796.
Okada, T. 1976. Subdivision of the genus Chymomyza Czerny (Diptera, Drosophilidae) with
description of three new species. Kontyu, Tokyo 44:496-511.
Schumann, H. 1987. Chymomyza amoena (Loew, 1862)-eine fUr die Fauna der DDR neue
amerikanische Drosophilidenart (Diptera). Entomologische Nachrichten und Berichte
31:125-127.
Sokal, R. R. and F. J. Rolhf. 1969. Biometry. second edition. W. H.Freeman, San Francisco.
859 pp.
50
Vol. 24, No.1
Spieth, H. T. 1987. The Drosophila fauna of a native California forest (Diptera: Drosophili
dae). Pan-Pacific EntomoJ. 63:247-255.
Sturtevant, A. H. 1921. The North American Species of Drosophila. Carnegie Inst. Wash.
PubJ. No. 301, Washington, D. C.
Throckmorton, L. H. 1962. The problem of phylogeny in the genus Drosophila. Univ. of
Texas PubJ. No. 6205:207-343.
Weckerly, F. W.• K. E. Nicholson and R. D. Semlitsch. 1989. Experimental test of discrimina
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Wheeler, M. R. 1952. The Drosophilidae of the Nearctic region exclusive of the genus Dro
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Williams, C. E. 1989a. Checklist of North American nut-infesting insects and host plants. 1.
Entomol. Sci. 24:550-562.
Williams, C. E. 1989b. Acorn Insects. Entomol. Notes. No. 20 ( 22 Nov). Mich. Entomol.
Soc.
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37:120-132.
1991
51
LIST OF THE LEPIDOPTERA OF BLACK STURGEON LAKE, NORTHWESTERN ONTARIO, AND DATES OF ADULT OCCURRENCE C.J. Sanders I
ABSTRACT
From May to September each year from 1960 through 1968, a collection of
Lepidoptera was made at Black Sturgeon Lake, northwestern Ontario, from speci
mens captured in a light trap and from specimens netted during the day. A total of
564 species was recorded from 70 families. A list of the species with dates of capture
is presented.
From 1960 through 1968, a 15-watt black-light trap was operated each year at a
Forestry Canada field station at Black Sturgeon Lake, northwestern Ontario. The
trapping program was initiated by R.E. Fye in 1960 and was taken over by C.J.
Sanders in 1965. Each year from 1960 through 1968, the light trap was set up when
the station was opened in early May and it was operated continually until the station
was closed in mid-to late September. In 1969 the research program at the Black
Sturgeon Lake field station was curtailed, and operation of the light trap was
discontinued.
Black Sturgeon Lake is located about 100 km from the northwestern shore of
Lake Superior, 110 km northeast of Thunder Bay at latitude 49"20'N, longitude
88°54'W, at an altitude of 250 m, in the Superior (B9) Boreal Forest Region of Rowe
(1972) (Fig. I). The corresponding Universal Transverse Mercator coordinates are
1636546. The physical, environmental and vegetational characteristics of the study
site have been described by Lethiecq and Regniere (1988). The site lies in the Black
Sturgeon District of the Lake Nipigon Site Region, which is characterized by: "Flat
topped basaltic ridges with deeper deposits-(i) granitic and clay forming sand, (ii)
silt, and (iii) limy clay, in the depressions." (Hills 1959). The vegetation is boreal
mixed wood, composed of fairly uniform stands of balsam fir (Abies balsamea),
white spruce (Picea glauca), black spruce (P.mariana), white birch (Betula papyri
feral, and trembling aspen (Populus tremuloides). Understory species include hazel
(Corylus cornuta), alders (both Alnus crispa and A. rugosa), and mountain maple
(Acer spicatum).
Continuous weather records are not available for the Black Sturgeon Lake area, so
instead information is presented for Cameron Falls, which is 42 km to the southeast of
Black Stugeon Lake (49°09'N, 88"21 'W, 229 m altitude), at the southern end of Lake
Nipigon (Anon. 1980). The mean annual air temperature is 1.7°C. The mean low
temperature for January is -16.6"C, with the lowest recorded temperature -46.1 "C.
The mean high for July is 17.0"C, with the highest temperature recorded 38.9°C. The
mean annual rainfall is 559.1 mm, and the snowfall 233.6 cm.
lForestry Canada, Ontario Region, P.O. Box 490, Sault Ste Marie, Ontario, Canada,
P6A SM7.
52
f2l
[I]]
D
~
•
Vol. 24, No. I
Boreal Predominantly Forest Forest and Barren Great Lakes - St. Lawrence Deciduous
Black Sturgeon Lake
Figure 1. Map of Ontario, showing location of Black Sturgeon Lake and boundaries of
Forest Regions (Rowe 1972).
The primary purpose of the light trap was to monitor the numbers of pest species
such as spruce budworm (Choristoneura jumijerana [Clem.]), jack pine budworm
(C. pinus pinus Free.) and forest tent caterpillar (Malacosoma disstria Hbn.), but
the opportunity was taken to assemble a collection representative of the Lepidoptera
in the area. The contents of the light trap were examined each morning, and repre
sentative specimens of each species were collected. Particular attention was paid to
obtaining the first specimens that were captured in the light trap each year. Addi
tional specimens of each species were collected thereafter throughout the flight
period, but no attempt was made to collect them in proportion to the numbers that
were collected in the trap. The collection is therefore an indication of the period over
which the insects were captured, and is not an indication of their abundance. In
addition to the light-trap material, specimens of day-flying Lepidoptera were col
lected by net.
All collected specimens were pinned and spread, and then sent for identification
to the Biosystematics Research Centre (Agriculture Canada, Ottawa, Ontario, Can
ada) or, after a reference collection had been assembled, identifications were made
by comparing specimens with the previously identified voucher specimens. Repre
1991
53
sentative specimens have been retained in the Canadian National Collection, in the
collection of the Forest Insect and Disease Survey (Forestry Canada, Ontario
Region, Sault Ste. Marie, Ontario) and in the personal collection of the author.
Insects in the following listing are catalogued according to Hodges et al. (1983).
Nomenclature has not been updated beyond that of Hodges et al. with the following
two exceptions: Anavitrinelia (checklist #6590) is incorrectly spelled by Hodges et al.
and has been corrected to Anavitrinella (e.g., McGuffin 1977). Cydia youngana
(Kft.) (checklist # 3466) is now considered to be a junior synonym of C. strobilella
(L.) (Brown and Miller 1983), and has been changed accordingly.
In several cases, entries at the subspecific level are accompanied by entries at the
corresponding specific level. In these instances, specimens under the species label
have not been identified to the subspecies level, and may therefore contain represen
tatives of one or more subspecies, including the accompanying subspecies that is
listed.
ANNOTATED LIST OF LEPIDOPTERA
HEPIALIDAE
COSMOPTERIGIDAE
19 Sihenopis purpurascens (Pack.) 15-30 Jul
20 Sthenopis quadrigultatus (GrL) 15 Jul
29 Hepia/us novigannus B. & Benj. 20 Aug-20
Sep
1515 Limnaeda phragmitella Staint. 20 Jul-IO
Aug
T1NEIDAE
261 Nemapogon acapnopennella (Clem.) 25
Jun-5 Jul
264 Nemapogon dejectella (Zell.) 15 Aug
421 Monopis spilotella Tengstrom 5 Jun-30 Jun
LYONETIIDAE
560 Buccu/atrix canadensisella Cham. 20 Jun
GRACILLARIIDAE
587 Ca/oplilia a/nivorella (Cham.) 15-30 May,
30 Jul-5 Aug, 20 Sep
601 Ca/optiJia coroniella (Clem.) 10-30 May
609 Ca/optilia invariabilis (Braun) 10 May
655 Parectopa pennsy/vaniella (Engel) 25 Jun
OECOPHORIDAE
911
913
914
916
Bibarramb/a allenella (Wism.) 30 Jun
Semioscopis merriccella Dyar 20 May-5 Jun
Semioscopis inornala Wlsm. 25 Aug
Semioscopis aurorella Dyar 20 May
BLASTOBASIDAE
1144 Gerdana caritella Bsk. 30 Jul-25 Aug
MOMPHIDAE
1458 Mompha um!asdella (Cham.) 30 Jun
GELECHIIDAE
1792 Co/eolechniles alrupictella (Dietz) 15-25
Jul
1826 Co/eolechnites piceaella (Kft.) 15 Jun-20
Aug
1869 Pseudole/phusa be/angerella (Cham.) 5-15
Jun
1923 Bryotropha clandeslina (Meyr.) 10-25 lui
1985 Gnorimoschema gallaeaslerella (Kellicott)
25 Jul-15 Aug
2069 Chionodes continuella (Zell.) 30 Jun-IO
luI, 5-15 Aug
2090 Chionodes /ugubrella (F.) 15 Jun-5 Jul
2093 Chionodes mediojuscella (Clem.) 30
May-20 Jun
2198 Aroga tria/bamacu/ella (Cham.) 15-20 Jun
2267 Brachmia jema/della (Bsk.) 10-30 Jun
2277 Dichomeris georgie//a (Wlk.) 20 May-IO
Jun
2302 Trichotaphe selosella Clem. 5-10 Aug
ALUClTIDAE
2313 A/ucita hexadacly/a L. 20-30 May, 5-20
Aug
PLUTELLIDAE
2366 Plulella xylostella (L.) 15-20 May
2376 Yps%pha dentijerella (Wlsm.) 25 JUI-25
Aug
2380 Ypsolopha falcijerella (Wlsm.) 20 Jul-S
Aug, 10 Sep
54
YPONOMEUTIDAE
2413 Swammerdamia caesiella (Hbn.) 15-20
Jun, 15 Jul
SESIIDAE
2554 Synan/hedon acerni (Clem.) 25 Jun-S Jul
CHOREUTIDAE
2651 Choreu/is diana (Hbn.) 20 Jul-IO Aug
COSSIDAE
2675 Acossus cen/erensis (Lin!.) 25 Jun-S Aug
2677 Acossus undosus (Lint.) 15-25 Jul
2693 Prionoxys/us robiniae (Peck) 5 Jul
TORTRICIDAE
Endo/henia a/bolineana (Kfl.) 30 Jun
Endo/henia impudens (Wlsm.) 20-30 Jul
Ap%mis funerea (MeYL) 5-25 Jul
Ap%mis spinu/ana (Me D.) 10-30 Jun
Apolomis /er/iana (McD.) 30 Jun
Apolomis dex/rana (McD.) 10 Jul-5 Aug
Pseudosciaphila duplex (Wlsm.) 10-15 Jul
Or/ho/aenia undu/ana (D. & S.) 5-25 Jul
O/e/hreu/es olivaceana (Fern.) 30 Jul
O/e/hreu/es zelleriana (Fern.) 30 JUI-15
Aug
2786 O/e/hreu/es punc/ana (Wlsm.) 25 Jul
2787 O/e/hreules connee/us (McD.) 10 Jul
2836 O/e/hreules cons/ella/ana (ZeU.) 20 Jul
2837 O/e/hreutes as/ra/ogana (Zell.) 15 Jul
2848 O/e/hreutes biparti/ana (Clem.) 25 Jul
2859 Olelhreu/es cespilana (Hbn.) 20 Aug
2892 Pe/rova albicapilana (Bsk.) 30 Jun
2898 Pe/rova gemislrigu/ana (KfL) 15 Jul
2900 Pelrova burkeana (Kft.) 15-30 Jul
2911 Phaneta awemeana (Kft.) 25-30 May
3116 Eucosma dorsisigna/ana (Clem.) 25 Aug-10
Sep
3116b.Eucosma dorsisignalana simi/ana (Clem.)
to Aug
3120 Eucosma dere/ec/a Heinr. 25 Aug
3211 N%ce/ia cu/minana (Wlsm.) 20 Jul-IO
Aug
3269 Grise/da radicana Heinr. 20 Aug-to Sep
3276a.Rhopobola unipunc/ana geminana
(Sleph.) 20-25 Jul
3280 Epinotia stroemiana (F.) 30 Aug-15 Sep
3283 Epino/ia so/andriana (L.) J 5 Jul-5 Aug
3286 Epino/ia medioviridana (KfL) 20 Aug-5
Sep
3302 Epinotia rec/iplicana (Wlsm.) 30 Jun-20
Jul
2745
2747
2755
2757
2759
2766
2769
2770
2778
2783
Vol. 24, No. 1
3304 Epinotia solici/ana (Wlk.) 30 May-25 Jun
3306 Epinotia nisella (CI.) 20-30 Jul, 10 Sep
3312 Epinotia momonana (KfL) 25 Jun-5 Jul,
15 Aug
3313 Epinotia ierracoctana (Wlsm.) to Aug
3347 Epinotia septemberana Kft. 30 Aug-15
Sep
3351 Epinolia lindana (Fern.) IS Aug-IO Sep
3355 Ancylis subaequana (Zell.) 15 Jun
3361 Aney/is semiovana (Zeit) 20 Jun
3380 Ancy/is goodelliana (Fern.) 30 May-20
Jun
3384 Ancy/is mediofasciana (Clem.) 5-25 Jun
3386 Ancy/is tineana (Hbn.) 15 Jun
3420 Pammene signijera (Heinr.) 20-30 Jun, 20
Jul
3428 Grapholila packard! Zell. 30 Jun
3466 Cyd!a strobilella (L.) to Jun, 5 Aug
3502 Croesia a/bicomana (Clem.) 20-25 Jul
3506 Ac/eris macdunnoughi Obr. 10 Sep
3508 Acleris ca/iginosana (Wlk.) 20 May. 20-25
Sep
3514 Acleris cervinana (Fern.) 20 May-IO Jun
3520 Acleris fuscana (B. & Bsk.) 2S May-15
Jun
3521 Acleris semiannu/a (Rob.) 15 May
3523 Acleris cornana (McD.) 10 lun
3525 Ac/eris forbesana (McD.) 20 Aug-25 Sep
3531 Acleris hastiana (L.) 15 May
3533 Acleris celiana (Rob.) 15 May-IO Jun
3540a.Acleris /ogiana p/acidana (Rob.) 30
Aug-25 Sep
3543 Acleris macu/idorsana (Clem.) 30 Aug-5
Sep
3548 Acleris variana (Fern.) 20 Jul-20 Aug, 30
Aug, 20 Sep
3549 Ac/eris maccana (fr.) 15 May
3551 Acleris inana (Rob.) 20-30 Aug
3556 Acleris nigrolinea (Rob.) 20 May-5 Jun, 30
Aug
3557 Acleris maximana (B. & Bsk.) 10 May
3565 Eulia ministrana (L.) 20 Jun-5 Jul
3595 Pandemis canadana Kft. 15 Jul-20 Aug
3625 Argyro/aenia mariana (Fern.) 5-10 lun
3632 Choristoneura fractivittana (Clem.) 5-10
Jul
3633 Choristoneura poralle/a (Rob.) 25 Jul
3635 Choristoneura rosaceana (Harr.) 20-30 Jul
3637 Choristoneura conf/ietana (Wlk.) 30
lun-IS Jul
3638 Choristoneura fumijerana (Clem.) 25
Jun-S Aug
3643 Choristoneura pinus Free. 25 Jul-S Aug
3648 Archips argyrospila (Wlk.) 25-30 Jul
3658 Arehips purpurana (Clem.) 5-15 Aug
1991
3661 Archips cerasivorana (Fitch) 30 Jul-IO
Aug
3664 Archips slriana (Fern.) 25 Jun-20 Jul
3665 Archips alberla (McD.) 10-30 Aug
3672 Syndemis affliclana (Wlk.) 20-25 Jun
3682 Clepsis persicana (Fitch) 5-20 Jul
3686 Clepsis melaleucana (Wlk.) 30 Jun
3688 Plycholoma perilana (Clem.) 20-25 Jul
3689 Plycholoma virescana (Clem.) 20-25 Jun
3697 Sparganolhis Iycopodiana (Kft.) 15 Aug
3706 Sparganolhis xanlhoides (Wlk.) 10-20 Jul
3720 Sparganolhis reliculalana (Clem.) 30 J ul-IO
Aug, 20 Sep
3740 Plalynola idaeusalis (Wlk.) 25 Jul
COCHYLIDAE
3787 Carolella vilellinana (Zell.) 25-30 Jun
3802 Hyslerosia riscana Kft. 15-20 Jul
HESPERIIDAE
3910 Thorybes pylades (Scudder) 10 Jul
3945 Erynnis icelus (Scudder & Burgess) 20 Jun
4041 Poliles Ihemislocles (Latr.) 20 Jul
4105 Amblyscirles vialis (Edw.) 20 Jun
PAPILIONIDAE
4176 Papilio glaucus L. 30 May-20 Jun
PIERIDAE
4195d.Arlogeia napi oleracea (Harr.) 15 Jun, 30
Jul-5 Aug
4197 Artogeia rapae (L.) 10 Jul
4210 Colias eurylheme Bdv. ]0 Jun, 20 Aug
4220 Colias inlerior Scudder 15 Jul
LYCAENIDAE
4362 Everes amynlula (Bdv.) 15-20 Jun
4372 Glaucopsyche Iygdamus (Doubleday) 15
Jun
NYMPHALIDAE
4423 Polygoniafaunus (Edw.) 5-25 Aug
4429 Polygonia progne (Cram.) 5 Aug, 25 Aug,
20 Sep
4430a.Nymphalis vau-album j-album (Bdv.&
Leconte) 5 Aug, 5-15 Sep
4432 Nymphalis anliopa (L.) 30 Jun, 5 Sep
4433 Aglais milberli (Ood!.) 5 Aug
4435 Vanessa cardui (L.) 25 May-5 Jun
4459 Speyeria allanlis (Edw.) 20-30 Jul
4464f.Clossiana selene alrocoslalis (Huard) 30
Jun, 30 Jul-5 Aug
55
4465a.Clossiana bellona loddi (Holl.) 20 Jun
4481 Phyctodes Iharos (Drury) 30 Jul-30 Jul
4490 Charidryas nycleis (Doubleday) 25 Jun
4522 Basilarchia arlhemis (Drury) 30 Jun-15 Jul
SATYRIDAE
4568 Enodia porllandia (F.) 15-20 Jul
LlMACODIDAE
4652 Torlricidia leslacea Pack. 30 Jun-25 Jul
PYRALIDAE
4703b.Gesneria cenluriella caecalis (Wlk.) 10-15
Jul
4716 Scoparia biplagialis Wlk. 30 Jun, 20 Jul-5
Aug
4737 Eudonia lugubralis (Wlk.) 15-20 Jun, 5
Jul
4748 Munroessa icciusalis (Wlk.) 15-30 Jul
4759 Parapoynx maculalis (Clem.) 10-20 Jul, 10
Aug
4760 Parapoynx obscuralis (Ort.) 20-25 Jul
4897 Evergeslis pallidala (Hufn.) 10-25 Jul
4935 Saucrobolys fumoferalis (Hulst) 20-30
Jun
4936 Saucrobolys fulilalis (Led.) 10-20 Jul
4950 Fumibolys fumalis (On.) 20-25 Jul, 25
Aug-IO Sep
4953a.Phlyclaenia coronala lerlialis (On.) 15-25
Jun
4956a.Nealgedonia eXlricalis dional (Wlk.)
20-25 Jun
4957 Muluuraia mysippusalis (Wlk.) 25-30 Jun
5060a.Pyrausla subsequalis borealis Pack. 10
Jun
5079 Udea rubigalis (On.) 30 May, 15 Jun, 30
Jun
5156 Nomophila nearclica Mun. 10 Jun-20 Jul,
20 Sep
5159 Desmia funeralis (Hbn.) 30 Jun-IO Jul
5262 Framinghamia helvalis (Wlk.) 25 Aug
5275 Herpelogramma perlexlalis (Led.) 10-25
Aug
5339 Crambus pascuellus (L.) 30 Jun-5 Jul, 20
Jul, 20 Aug
534 I a.Crambus alienellus labradoriensis
Christoph 10 Jul
5343a.Crambus perlellus innolalellus Wlk .. 15
Jul-5 Aug
5344 Crambus unislrialellus Pack. 25 Jul-IO
Aug
5357 Crambus leachellus (Zinck.) 10 Aug-5 Sep
5362 Cram bus agilalellus Clem. 20 Jul
5379 Crambus luteolellus Clem. 15-30 Jun
56
5380 Crambus zeellus Fern. 20 Jul
5391 Chrysoteuchia top/aria (Zell.) 30 Jun-20
Jul
5403 Agriphifa vulgivagel/a (Clem.) 10-20 Aug
5408 Catoptria lattradiella (Wlk.) 25 Jul-15
Aug
5413 Pediasia trisecta (Wile.) 10-20 Aug
5517 Aglossa caprealis (Hbn.) \5-30 Jul
5529 Herculia thymetusalis (WIle.) 30 Jun-15
lui
5655 Acrobasis tricolorella Grt. 15-30 Jun
5688 Acrobasis betulella Hulst 30 Jun-IO Aug
5716 Myelopsis coniella (Rag.) 10 Aug
5718 Myelopsis subtelricella (Rag.) 5-20 lun
5721 Apomyelois bistriatella (Hulst) 15 Aug, 10
Sep
5745 Glyptocera consobrinella (Zell.) 25 Jun
5798 Nephoptertx carneella (Hulst) 20-30 Jun
5812 Telethusia ovalis (Pack.) 15 Aug
5843 Dioryctria reniculelloides Mutuura & Mun.
101un, 10-15 Jul, IS Aug
5999 Eulogia ochrifrontella (Zell.) 30 Jun· JO
Jul, 15 Aug
6053 Peoria approximella (Wlk.) 5 lui
PTEROPHORIDAE
6102 Trichopti/us lobidactylus (Fitch) 20-25 lui
6157 Adaina montana (Wlsm.) 20 Aug
6213 Oidaematophorus lacteodactylus (Cham.)
10 Jun
THYATIRIDAE
6235 Habrosyne scripta (Gosse) 20-30 Jun, 20
Jul
6237 Pseudothyatira cymatophoroides (Gn.)
20-25 Jun, 5-30 Jul
6240 Euthyatira pudens (Gn.) 20 May-5 Jun
DREPANIDAE
6251 Drepana arcuata Wlk. 5 Jun-IO Jul
6252 Drepana bilineata (Pack.) 20-25 May,
10-15 Aug
6255 Orela rosea (Wlk.) 10 lul-IO Aug
GEOMETRIDAE
6256 Archiearis injans (Masch.) 20 May
6279 ftame occiduaria (Pacle.) 25 Jul
6286 /tame brunneata (Thunb.) 10-15 Jul
6304 ftame bitactata (WJIe.) 15 Jul·20 Aug
6330 Semiothisa ulsterata (Pears.) 30 Jun-20
Jul
6352 Semiothisa granitata (Gn.) 30 Jun
6373 Semiothisa denticulata Grt. 25 Jun
6394 Semiothisa hebetata (Hulst) 10 Jul
Vol. 24, No.1
6428 Orthojidonia tinctaria (Wlk.) 5-10 May, 15
Jun
6570 Aethalura intertexta (Wlk.) 10-15 Jun
6583 Anacamptodes ephyraria (WIle.) 15-20
Aug
6588 fridopsis larvaria (Gn.) 25 Jun
6590 Anavitrinella pampinada (Gn.) 30 Jun
6597 Ectropis crepuscularia (D. & S.) 25 May-S
Jun
6621 MelaTlolophia signataria (Wlk.) 5 Jun
6640a.Biston betularia cognataria (Gn.) 10-25
Aug
6666 Lomographa semiclarata (Wlk.) 20-25
May
6668 Lomographa glomeraria (Ort.) 20-30 May
6678 Cabera variolaria On. 25 Jun-20 Jul
6726 Euchlaena obtusaria (Hbn.) 30 Jun-5 Jul
6731 Euchlaena madusaria (Wlk.) 5-20 Jul
6734 Euchlaena marginaria (Minot) 30 May-15
Jun
6740 Xanthotype urticaria Swett 30 Jun
6743 Xanthotype sospeta (Drury) 30 Jun-20 Jul
6754 Pero hubneraria (Gn.) 5-30 Jun
6796 Campaea per/ata (On.) 10-25 Jui
6797 Ennomos magnaria Gn. 10-25 Aug
6807 Tacparia detersata (Gn.) 25-30 May
6818 Se/enia kentaria (G. & R.) 20 May-IO Jun
6819 Metanema inatomada Gn. 30 May, 15
Jun-5 Jul
6820 Metanema determinata Wlk. 15 Jun-20
Jul
6822 Melarran/his duaria (On.) 25 May-5 Jun
6836 Anagoga occidu(1ria (Wlk.) 25 May-20
Jun
6838 Probole amicaria (H.-S.) 25 Jun-15 Jul
6841 Plagodis kuetzingi (Grt.) 30 Jun
6842 Plagodis phlogosaria (Gn.) 30 MaY-IS Jun
6863 Caripeta divisata Wlk, 20 Jun-IO Jul
6885 Besma quercivoraria (On.) 20 Jun
6888 Lambdina jiscellaria (Gn.) 30 Aug-25 Sep
6906 Nepytia canosaria (Wlk.) 25 Aug-20 Sep
6912 Sicya macularia (Harr.) 10-30 Aug
6964 Tetracis cachexia/a On. 25-30 lun
6982 Prochoerodes transversata (Drury) 10
Aug-20 Sep
7009 Nematocampa limbata (Haw.) 10-15 Aug
7048 Nemoria mimosaria (Gn.) 20 lun
7058 Synchlora oerata (F.) 5-25 Jul
7139 Cyclophora pendulinaria (On,) 15-30 Jun,
20 Jul
7159 Scopula limboundata (Haw.) 15 Jul
7162 Scopula ancel/ata (Hulst) 25 Jul
7164 Scopula junctaria (Wlk.) 10 Jul
7182 Dysslroma citrata (L.) 15-25 Aug, 15 Sep
7187a.Dysstroma truncata traversal (Kellicott) 5
Jul
1991
7196 Eulilhis diversilineata (Hbn.) 20-25 Aug
7201 Eulithis testata (L.) 25 Aug-IO Sep
7203 EuHthis molliculala (Wlk.) 5-15 Aug
7206 Eulilhis explanata (Wlk.) 20 Jul-IS Aug
72l3a.Ecliplopera silaceata albolineal (Pack.) 15
Jun·30 Jul
7216 P/emyria georgii Hulst 20 Aug, 20 Sep
7239 Hydriomena pluviata (Gn.) 30 Jun
7254 Hydriomena ruberata (Freyer) 30 May-IS
Jun
7257 Hydriomena furcata (Thunb.) 5 Sep
7285 Triphosa haesitata (Gn.) 10-25 May
7285a.Triphosa haesitata affirmari (Wlk.) 20-30
May, 5-25 Sep
7291 Hydria undu/ata (L.) 30 Jul
7293 Rheumaptera hastata (L.) 5-20 Jun
7294 Rheumaptero subhastata (Nolcken) 10-25
Jun
7307 Mesoleuca rujicillala (Gn.) 30 Jun
7312 Spargania magnoliata Gn. 30 Jun. 25 Jul
7313 Spargania lucluala (D. & S.) 30 Jun
7313a.Spargania lucluala obducta (Mtisch.) 30
Jun-IS Aug
7316 Perizoma basaliata (Wlk.) 10-15 Aug
7329 Anticlea vasiliata Gn. 15 May-5 Jun
7330 Antic/ea multiferata (Wlk.) 20-25 Jun
7368 Xanthorhoe labradorensis (Pack.) 10 Jul
7384 Xanthorhoe munltata (Hbn.) 30 Aug
7388 Xanthorhoe ferrugata (CL) 5-30 Jun
7390 Xanthorhoe lacustrata (Gn.) 15 Jun
7394 Epirrhoe alternata (Muller) 5 Jul. 25 Jul
7414 Orthonama obstipata (F.) 20 May
7416 Orthonama centrostrigaria (Woll.) 25 Jun
7422 Hydre/ia inornata (Hulst) 10 Jul
7425 Venusia cambrica (Cur!.) 20-30 Jul, 30
Aug
7428 Venusia comptaria (Wlk.) 30 MaY-5 Jun
1445 Horisme intestinata (Gn.) 15-25 Jun
7459 Euplthecia co/umbiata (Dyar) 20-25 May
7416 Eupithecia misturata (Hulst) 5 Jun
7481 Eupithecia coloradensis (Hulst) 20-25 Jun
7522 Eupithecia nimbicolor (Hulst) 25 Jun
7526 Eupithecia russeliata Swett 10-15 Jun
7529 Eupithicia coagufata Gn. 20 May·25 Jun.
10-20 Aug
7543 Eupithecia annu/ata (Hulst) 25-30 May
7575 Eupithecia mutata Pears. 20 Jun
7594 Eupithecia anticaria Wlk. 30 Jun
7605 Eupithecia ravocostafiata Pack. 25-30
May
7635 Acas!s viridata (Pack.) 30 May·5 Jun
7637 Cladara limitaria (Wlk.) 15 May·5 Jun
7639 C/adara atroliturata (Wlk.) 20 May-5 Jun
57
EPIPLEMlDAE
7650 Callizzia amorata Pack. 30 Jun-20 Jul
LAS10CAMPlDAE
7673 Tolype lariels (Fitch) 25 Aug-IO Sep
7687 Phyl/odesma americana (Harr.) 15 May-5
Jul
7698 Mafacosoma disstria Hbn. 5 Aug
7702c.Malacosoma californicum pluvial (Dyar)
30 Jul
SATURNIIDAE
7757 Anlheraea polyphemus (Cram.) 20-25 Jun
7758 Actias luna (L.) 20 Jun·1O Jul
SPHINGIDAE
7787
7809
7810
7811
7817
7821
Ceratomia undulosa (Wlk.) 30 Jun·30 Jul
Sphinx kalmiae J. E. Smith 15 Aug
Sphinx gardius Cram. 20-30 Jun
Sphinx luscitiosa Clem. 30 Jun
Lapara bombycoides Wlk. 10-15 Jul
Smerinthus jamaieensis (Drury) 5 Jun. 20
Jun-5 Jul, 15-20 Jul
7822 Smer!nthus eerys! Kby. 5-20 Jun. 5 Jul
7824 Paonias excaecatus (J. E. Smith) 10-25
Jun
7827 Laothoe jug/andis (J. E. Smith) 30 May-5
Jul
7828 Pachysphinx modesta (Han.) 25-30 Jun, 5
Sep
7854 Hemaris gracilis (G. & R.) 15-25 Jun
1855 Hemaris diffinis (Bdv.) 25 Jun
1893 Hyles gallii (Rottemburg) 5-15 Jul, 5 Aug
NOTODONTIDAE
7895 Clostera albosigma Fitch 25 May-15 Jun.
15 Jul, 5 Aug
7915 Nadata gibbosa (1. E. Smith) 10 Jun·1O
Jul, 20 Jul
1921 Peridea ferruginea (Pack.) 15 Jun-30 Jul
7922 Pheosia rimosa Pack. 15 Jul, 10 Aug
7924 Odontosia elegans (Stkr.) 20 Jun-IO Jul. 25
Aug
7926 Notodonta scitipennis Wlk. 10-15 Jul
1928 Notodonta simp/aria Graef 30 May-30
Jun, 15 Jul, 10 Aug
7934 Gluphisia lintneri (Grt.) 20 May-15 Jun
7939 Furcula occidenta/is (Lin!.) 5-10 Jun. 5
Jul
7941 Furcula modesta (Hudson) 30 May·20 Jul
7990 Heterocampa umbrata Wlk. 25 Jun-15 Jul
7990a.Heterocampa umbrata pulverea (G. & R.)
20 Jun·1O Jul
58
8005 Schizura ipomoeae Doubleday 5-20 Jul
8007 Schizura unicornis (J. E. Smith) 15 Jul·;
Aug
8011 Schizura leplinoides (Grt.) 20 Jun-25 Aug
ARCTIIDAE
8045.1 Crambidia pallida Pack. 25 Aug
8051 Crambidia casta (Pack.) 20 Aug·1O Sep
8090 Hypoprepia /ucosa Hbn. 10 Aug
8098 Clemensia albata Pack. 10-25 Aug
8111 Haploa lecontei (GUi:r.-Meneville) 30
Jun·20 Jul
8114a.Holomelina lae/a treatii (Grt.) 25 Jun·15
Jul
8123 Holomelina /erruginosa (Wlk.) 25 Jun·20
Jul, 5 Aug
8127 Parasemia plan/aginis (L.) 25 Jun.1O Jul
8134 Spilosoma congrua Wit. 5-20 Jun
8136 Spilosoma dubio (Wlk.) 20 Jun
8137 pilosoma virginica (F.) 20-30 Jun
8158 Phragmafobia assimilans Wlk. 25 May·1O
Jun
8162 Pla/arc/ia parthenos (Harr.) 20 Jun·; Jul
8166 Arc/ia caja (L.) 15-30 Aug
8175 Apantesis virguncula (W. Kirby) 25-30 Jul
8176 Apantesis anna (Grt.) 25 Jul
8186 Apantesis williamsii (Dodge) 5-15 Jul
8187 Apantesis celia (Saund.) 20 Jul
8196 Apanlesis par/henice (W. Kirby) 15 Jul-20
Aug
8197 Apan/esis virgo (L.) 15 Jul
8214 Lophocampa macula/a Harr. 20-30 lun
8262 C/enucha virginica (Esp.) 10 Jul-5 Aug
8267 Cisseps /ulvico/lis (Hbn.) 10-20 Jul
NOCTUIDAE
8322 [dia americalis (Gn.) 25 Jul, 10 Aug
8334 [dia fubricalis (Gey.) 25 J ul-IO Aug
8338 Phalaenophana pyramusalis (Wlk.) 10 lun
8341 Zanclognalha lheralis (Wit.) 5-20 Jul
8349 Zanclognatha protumnusalis (Wit.) 5 Jul
8352 Zancfogna/ha jaeehusalis (Wit.) 15 Jul
8356 Chytolila pe/realis Grt. 20-30 lun
8362 Phalaenoslola melonalis (Wlk.) 5-25 Jul
8370 Bleptina earadrinalis Gn. 10 Jul, 20 Aug
8397 Palthis angulalis (Hbn.) 15-25 Jun
8413 Mye/erophora inexplica/a (Wlk.) 10-20
lui
8443 Bomolocha bijugalis (Wlk.) 25 Jul
8444 Bomolocha palparia (Wlk.) 5 Jul
8455 Lomanaltes eduetalis (Wlk.) 25 Jul
8461 Hypena humuli Harr. 20 Aug
8479 Spargaloma sexpunctata Grt. 30 Jun-IO
Jul, 5 Aug
8490 Pangrap/a decoralis Hbn. 30 Jun
Vol. 24, No. 1
8500 Metalectra quadrisigna/a (Wlk.) 30 lun·1O
Jul, 25 Jul
8554 Alabama argillacea (Hbn_) 5 Sep
8694 Zale aeruginosa (Gn.) 25 May-15 Jun
8697 Zale minerea (On.) 5-15 lun
8697a.Zale minerea norda (Sm.) 25 Jun·5 Jul
8703 Zale duplicata (Bethune) 10-15 lun
8716 Zale unilinea/a (Grt.) 25 May-5 Jun
8731 Eucfidia cuspidea (Hbn.) 10 Jun-5 Jul
8738 Caenurgina crassiuscula (Haw.) 25 Jun, 30
Jul, 15 Aug
8803 Catocala relicta Wlk. 20 Aug-5 Sep
8805 Catocala unijuga Wlk. 5-20 Aug
8817 Catocala briseis Edw. 20 Aug-20 Sep
8821 Catocala semirelicta Grt. 20 Aug
8833 Calocala concumbens Wlk. 30 Aug-IO Sep
8846 Catocala sordida Grt. ; Aug
8857 Co/ocala ultronia (Hbn.) 20 Aug·15 Sep
8867 Catocala blandula Hulst 30 Jul-25 Aug
8896 Diachrysia aero ides (Grt.) 20 Jul·1O Aug
8897 Diachrysia balluca Gey. 25 Jul-15 Aug
8899 Pseudeva purpurigera (Wlk.) 30 Jul-IO
Aug, 25 Aug
8904 Chrysanympha formosa (Ort.) 5 lul·15
Aug
8908 Autographa precationis (Gn.) 15 Jun,
15-20 Jul, 25 Aug, 20 Sep
8909 Au/ographa rubida Otto!. 15 Jun·5 Jul
8911 Autographa bimaeula/a (Steph.) 25 Jul·25
Aug
8912 Autographa mappa (G. & R.) 30 Jun-5
Aug
8913 Autographa pseudogamma (Ort.) 5-15 JuI
8916 Autographa /Iagellum (Wlk.) 25-30 Jul, 5
Sep
8923 Autographa ampla (Wlk.) 15 Jul·15 Aug
8924 Anagrapha /ald/era (Kby.) 25 May, 15-30
lun, 25 Aug-IO Sep
8926 Syngrapha octoscripta (Grt.) 10 Jul, 5-25
Aug
8927 Syngrapha epigaea (GrL) 15-25 Aug
8928 Syngrapha selee/a (WIk.) 15 Jul·20 Aug
8939 Syngrapha alias (Ottol.) 25 Jun, 10 Aug
8942 Syngrapha rec/angula (W. Kirby) 5 JuI, 25
Jul, 15-25 Aug
8945 Syngrapha montana (Pack.) 30 Jun
8946a.Syngrapha microgamma neare/ica Fgn. 30
Jun
8950 Plusia putnami Grt. 30 Jun.1O Sep
8953 Plusia venusta Wlk. 30 Jul-5 Aug
8969 Baileya doubledayi (Gn.) 30 lun
8970 Baileya ophthalmica (Gn.) 5 Jun
8974 Characoma nilofica (Rogenhofer) 25 Jun,
30 Jul
9046 Li/haeodia bellicula Hbn. 10 Jut
9048 Lithaeodia albidula (Gn.) 20-25 Jun
1991
9053
9059
9090
9177
Lithacodia carneola (Gn.) 30 Jun-IS Jul
Capis curvata Grt. 10-20 Jul
Tarachidia candefacta (Hbn.) 15 Sep
Panthea acronyctoides (Wlk.) 20 Jun-5
Jul
9183 Panthea pallescens McD. 30 Jun, 25-30
Jul
9183a.Panlhea pallescens centralis McD. IS Jul
9185 Colocasia propinquilinea (Grt.) 30 May-15
Jun, S Jul
9189 Charadra deridens (Gn.) 20 Jun, 15-20
lui
9193 Raphia frater Grt. 15-20 Jun
9203 Acronlcta dactylina Grt. 20-30 Jun, 5 Sep
9205 Acronicta lepusculina Gn. 20 lun
9207 Acronlcta innotata Gn. 5-25 Jun, 10-20
Jul, 10-20 Aug
9212 Acronlcta grisea Wlk. 20 lun-IO Jul
9226 Acronlcta superans Gn. 25-30 Jun
9229 Acronicta hasta Gn. 25 Jun
9241 Acronicta frag/lis (Gn.) 20-25 Jun, 10-15
Jul, 15 Aug
9257 Acronlcta impleta Wlk. 20 Jun
9259 Acronlcta noctivaga Grt. 10-20 lun
9286 Harrlsimemna trisignata (Wlk.) 30 Jun-5
Jul
9326 Apamea verbascoldes (Gn.) 20 Jul-S Sep
9344 Apamea plutonla (Grt.) 25 Jul-5 Aug
9351 Apamea alia (Gn.) 25 Jun. IS lui
9359 Apamea commoda (Wlk.) 5-30 Jul
9360 Apamea impulsa (Gn.) 15 Jul
9365 Agroperlna lateritia (Hufn.) 30 Jun-IO Jul
9367 Agroperlna dubitans (Wlk.) 20 Jul, 20
Aug
9367a.Agroperlna dubilans cogitata (Sm.) 10-15
Jul, 30 lui, 25 Aug
9374 Protagrotis nivelvenosa (Grt.) 5-10 Aug
9382 Crymodes devastator (Brace) 10 Aug-5
Sep
9391 Luperina passer (Gn.) 10 Aug
9415 Oligia bridghaml (G. & R.) 25 Aug
9419 Oligla mactala (Gn.) 20 Jul, 20 Aug-15
Sep
9420 Oligia iIlocata (Wlk.) 20-30 Jul, 25 Aug
9431 Parastichtis disc/varia (Wlk.) IS Aug-5
Sep
9437 Hypocoena Inquinata (Gn.) 25 Aug
9439 Hypocoena basistr/ga (McD.) 5-20 Sep
9449 Archanara oblonga (Grt.) 5-10 Sep
9450 Archanara subflava (Grt.) 5 Sep
9453 Helotropha reniformis (Grt.) 5-20 Sep
9454 Amphipoea velata (Wlk.) 20 Aug
9457 Amphipoea americana (Speyer) 5 Aug-lO
Sep
9472 Papalpema harrisli (Grt.) 20 Aug·20 Sep
9474 Papaipema sauzalitae (GrL) 30 Aug-5 Sep
9480
9509
9520
9523
59
Papa/pema pterlsii Bird 20 Sep
Papaipema unlmoda (Sm.) 5-25 Sep
Achatodes zeae (Harr.) 30 Jul. 25 Aug
Bellura gortynoldes f. diffusa (Grt.) 25
lun
9525 Bel/ura obliqua (Wlk.) 20 lun-20 luI
9545 Euplexia benesimills McD. 25-30 Jun
9546 Phlogophora 'ris Gn. 20-30 Jun
9547 Phlogophora periculosa Gn. 30 lul-25
Aug
9549 Enargia decolor (Wlk.) 15 Aug·20 Sep
9555 lplmorpha pleonectusa Grt. 15 Aug.20
Sep
9556 Chytonix palliatrlcula (Gn.) 20 Jun-15 Jul,
15-25 Aug
9560 Dypterygia rozmani Berio 10-20 Jul
9564 Andropo/la contacta (Wlk.) 15-30 Aug
9578 Hyppa xylinoides (Gn.) 20 Jun·20 Jul. S
Sep
9633 Callopistria cordata (Ljungh) 25 Jun-20
Jul
9637 Magusa orbifera (Wlk.) 10 Sep
9647 Proxenus miranda (Ort.) 20-30 Jun
9649 Proxenus mendosa McD. 20 Jul
9657 Platyperlgea multifera (Wlk.) 10 Aug· I 0
Sep
9666 Spodopterafrugiperda (J. E. Smith) 10-20
Sep
9681 Elaphriafestlvoides (On.) 5 Jun-IO Jul
9873 Xylena nupera (Lint.) 5 May
9874 Xylena curvimacula (Morr.) 20 May-5 Jun.
20 Sep
9876 Xylena cineritia (Ort.) 10-25 May. 30
Aug-20 Sep
9878a.Lithomoia solidaginis germana (Morr.) 20
Aug·25 Sep
9884 Litholomia napaea (Morr.) 10-25 May.
10-25 Sep
9889 Llthophane petulco Grt. 20-25 May. 20
Sep
9891 Lithophane amanda (Sm.) 5 Jun
9902 Lithophane baileyi Grt. 15-25 Sep
9913 Lithophane georgii Grt. 10 May
9916 Llthophane unimoda (Lint.) 15 May-5 Jun,
25 Sep
9917 Llthophane fagina Morr. 10 May-IO lun.
25 Sep
9922 Lithophane pexata Grt. 10-20 May. 10-25
Sep
9928 Lithophone thaxteri Grt. 10-20 May, 20
Sep
9935 Eupsi/ia trlstigmata (Grt.) 10 May, 20 Sep
9939 Eupsi/ia devia (Grt.) 15-20 May
9952 Eucirroedia pampina (On.) 15 Aug-15 Sep
9961 Anathix ralla (G. & R.) 5 Sep
9962 A nathix pula (G. & R.) 15 Aug, 5-20 Sep
60
Xanthia logata (Esp.) 30 Aug-IO Sep
Hillia iris (Zett.) 5-15 Sep
Fi~hia en thea Grt. 10-25 Sep
Plalypolia anceps (Steph.) 5-20 Sep
Xylotype acadia B. & Ben]. 30 Aug-25
Sep
9990 Sulyna profunda (Sm.) 25 Aug-20 Sep
9998 Brachylomia algens (Grt.) 10 Sep
10005 Feralia jocosa (Gn.) 15 May-IO Jun
10007 Feralia major Sm. 15-20 May, 5 Jun
10008 Feralia comstock! (Grt.) 25 May-IO Jun
lDOIl Brachionycha borealis (Sm.) 5 May-30
Jun
10059 Homohadena badislriga (Grt.) 30 Jul-30
Aug
10065 Homohadena infixa (Wlk.) 10 Jul-20
Aug
10123 Oncocnemis piffardi (Wlk.) 10 Sep
10198 Cucutlia postera Gn. IS J ul
10223 Disceslra trifolii (Hufn.) 10 Sep
10265 Sideridis rosea (Harv.) 10-20 Jun
10275 Polia nimbosa (Gn.) 20 lul-IO Aug
10276 Polia imbrifera (Gn.) 20 JuI-5 Aug
10280 Polia purpurissata (Grt.) 15 JuI-20 Aug
10288 Polia detracta (Wlk.) 10 Jul
10288a.Polia detracta neoterica (Sm.) 15-25 lui
10290 Polia obscura (Sm.) 15-25 Jun
10292 Melanchra adjuncta (Gn.) 10-25 Jun.
10-25 Jul
10294 Melanchra pulverulenta (Sm.) 15-30 Jun
10295 Melanchra assimilis (Morr.) 30 Jun-15
Jul
10296 Lacanobia nevadae (GrL) 10 Jun-5 Jul.
15 Aug
10298 Lacanobia radix (Wlk.) 5-15 Jun
10300 Lacanobia grandis (Gn.) 5 Aug
10301 Lacanobia lutra (Gn.) 10-25 Jun
10303 Lacanobia tacoma (Stkr.) 10-25 Jun
10310 Papestra quadrala (Sm.) 30 May-5 Jun
10312 Papestra cristifera (Wlk.) 5-30 Jun
10370 Lacinipolia lustra/is (GrL) 20 Jun-IO Jul
10372 Lacinipolia anguina (GrL) 5-15 Jun
10397 Lacinipolia renigera (Steph.) 25 Jul-30
Aug
10405 Lacinipolia lorea (Gn.) 30 Jun·1O Jul
10436 A/etia oxygala (GrL) 20 Jul. 10-25 Aug
10438 Pseudaletia unipuncta (Haw.) 10 Jun
10446 Leucania mu/tilinea Wile. 10-20 Aug
10447 Leucania commoides Gn. 5-30 Jun
10449 Leucania insueta Gn. 15 Jun
10471 Slretchia plusiaeformis Hy. Edw. 15-20
May
1047 I a.Stretchia p. coloradico/a Strand. 5-20
May
10490 Orthosia revicta (Morr.) 10-25 May
10513 Egira d%sa (Grt.) 25 May·IO Jun
9965
9967
9974
9976
9980
Vol. 24, No. 1
10563 Protorthodes oviduca (Gn.) 10-25 Jun
10578 Pseudorthodes vecors (Gn.) 30 Jun·5 Jul
10587 Orlhodes cynica Gn. 10 Jun. 15 Jul
10644 Agrotis mollis Wlk. 25 Jul
10651 Agrolis venerabilis Wlk, 20 Aug-15 Sep
10659 Agrolis volubilis Harv. 25 Jun
10660 Agrotis obliqua (Sm.) 15 Jul
10663 Agrolis ipsi/on (Hufn.) S-25 Sep
10670 Fe/tia jaculifera (Gn,) 30 Aug·15 Sep
10676 Feltia herilis (GrL) 2S Jul·1O Sep
10702 Euxoa divergens (Wlk.) 30 Jun-30 Jul
10715 Euxoa scandens (Riley) 25-30 Jul. 10-15
Sep
10738 Euxoa mimallonis (GrL) 5 Aug-S Sep
10755 Euxoa declarata (Wlk.) 5-25 Aug
10801 Euxoa ochrogaster (Gn.) 5-15 Aug
10891 Ochropleura plecta (L.) 5-15 Jul, 25 Aug
10915 Peridroma sauda (Hbn.) 15 May-20 Jun.
5-25 Sep
10917 Diarsia rubifera (GrL) 30 Jul·20 Aug
10918 Diarsia dis/ocata (Sm.) 30 Jul·15 Aug
10919 Diarsia jucunda (Wlk.) 15 Jul·1O Sep
10928 Graphiphora haruspica (GrL) 30 Jul-25
Aug
10929 Eurois occulta (L.) 5 Jul, 10-25 Aug
10930 Eurois astricta MorT. 20 Jul·25 Aug
10942 Xestia adela Franc. 5 Aug·1O Sep
10943 Xestia normaniana (Gr!.) 20 Jul·20 Aug
10944 Xeslfa smithii (Snell.) 20 Jul, 10-20 Aug
10947 Xestia ob/ata (Morr.) 5 Jul·1O Aug
10951 Xestia tenuicu/a (Morr.) 5 Aug
10954 Xestfa col/oris (G. & R.) 25 Aug
10957 Anomogyna atra/a (Morr.) 25-30 Jun
10962 Anomogyna perquiritata (Morr.) 10-30
Aug
10967 Anomogyna elimata (Gn.) 10 Aug-1O Sep
10969 Anomogyna dilucida (Morr.) 5 Aug
10988a.Eugraphe subrosea opacifrons (Grt.) 20
Aug
10992 Paradiarsia tit/oratis (Pack.) 5-15 Jul
10996 Meta/epsis saticarum (Wlk.) 15-25 May
10997 Meta/epsis fishii (Grt.) 30 Jun
10999 Aplectoides colUiita (Gn.) 20-25 Jun, 20
Jul
11000 Anaplecloides prasina (D. & S.) 30 Jun·5
Jul, 15 Jul. 15 Aug-15 Sep
11001 Anaplectoides pressus (Grt.) 5 Jul·1O Sep
11003 Chersotisjuncta (Grt.) 25 Jul-IS Aug
11004 Proto/ampra rufipectus (Morr,) 5-25
Aug
11008 Eueretagrolis peraltenta (GrL) 5 Jul
1991
11009 Eueretagrotis atlenta (Grt.) 30 Jun-20
Aug
11010 Heptagrotis phyllophora (Grt.) 25 Jul
11012 Cryptocala acadiensis (Bethune) 10 Jul-IS
Aug
61
11043 Rhynchagrotis cupida (Grt.) 15 Sep
11064 pyrrhia exprimens (Wlk.) 25 Jun-S Jul
11068 Heliothis zea (Boddie) IS-20 Sep
DISCUSSION
Black Sturgeon Lake is in a relatively remote area, where only limited collecting
has been carried out in the past. In his review of the state of knowledge on the
Lepidoptera in Canada, Munroe (1979) lists a number of locations from which
faunal samples are needed. Among these he names "the Lakehead district of
Ontario, and the whole region from there to the Lake of the Woods, along the U.S.
border." Black Sturgeon Lake is situated at the eastern end of this area, and this
listing is therefore a contribution. to meeting this lack of knowledge.
In all, 564 species of Lepidoptera plus 8 subspecies were collected at Black Stur
geon Lake during the years 1960-1968, representing 36 families, 5 butterflies, 1
skipper and 30 moths. This compares with 4692 species from 70 families listed by
Munroe (1979) for the whole of Canada. It is probable that the listing for Black
Sturgeon Lake is incomplete. The collections were made by staff trained in forest
entomology, but with no' taxonomic expertise, who may have overlooked rarer
species, especially if they were inconspicuous or resembled other, more numerous,
species. However, it is unlikely that many species were overlooked. The stated
objective was to make a representative collection, and so the collectors were contin
ually on the look out for new species. The listing may then be considered fairly
reliable, but absence from the listing cannot be taken as conclusive evidence that the
species is absent from the area.
ACKNOWLEDGMENTS
I thank members of the Biosystematics Research Center (Agriculture Canada) for
the identification of numerous specimens, and all those at Forestry Canada, Ontario
Region, who worked on the assembly, typing and proofreading of the information
in this report. In particular, Doug Lawrence, George Lucuik, Suzanne Palmer,
Elizabeth Chudoba and Corrine Motluk should be thanked; without their care,
diligence, and patience, the work would never have been completed. As well, my
thanks to Dr. Paul Syme (Forestry Canada, Ontario Region) and Dr. Kevin Barber
(Forestry Canada, Forest Pest Management Institute) for their careful and thorough
review and for their advice on the taxonomy and nomenclature of many of the
species and subspecies.
LITERATURE CITED
Anon. 1980. Canadian climate normals, 1950-1980: temperature and precipitation. Dep. Envi
ron., Atmospher. Environ. Serv., Downsview, OnL, 254 pp.
Brown, R.L., and W.E. Miller. 1983. Valid names of the spruce seed moth and a related Cydia
species (Lepidoptera: Tortricidae). Ann. entomol. Soc. Amer. 76:110-111.
Hills, O.A. 1959. A ready reference to the description of the land of Ontario and its productiv
ity. Preliminary Report. Onto Dep. Lands and Forests, Div. Res. 142 pp.
Hodges, R.W., T. Dominick, D.R. Davis, D.C. Ferguson, J.O. Franclemont, E.O. Munroe
and J .A. Powell. 1983. Check List of the Lepidoptera of America North of Mexico. E. W.
Classey Ltd., Faringdon, Oxfordshire, England, and The Wedge Entomological Research
Foundation, Washington, D.C., U.S.A., 284 pp.
62
Vol. 24, No. 1
Lethiecq, L-L., and J. Regniere. 1988. Comparative study of six sites used by the Canadian
Forestry Service in the study of spruce budworm population dynamics. Gov't. of Can., Can.
For. Serv., Ste.-Foy, Quebec. Inf. Rep. LAU-X-83. 46 pp.
McGuffin, W.C. 1977. Guide to the Geometridae of Canada (Lepidoptera) II. Subfamily
Ennominae. Mem. ent. Soc. Can. 101, 191 pp.
Munroe, E. 1979. Status of taxa in Canada: Lepidoptera. in H.V. Danks (ed.), Canada and its
Insect Fauna. Mem. ent. Soc. Can., No. 108, pp. 427-481.
Rowe, 1.S. 1972. Forest Regions of Can., Dep. Environ., Can. For. Serv., Ottawa, Onto Pub!.
No. 1300. 172 pp.
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unable to pay costs from personal funds, may apply to the Society for financial assistance. Application for
subsidy must be made at the time a manuscript is initially submitted for publication.
Authors will receive page proof, together with an order blank for separates. Extensive changes to the
proof by the author will be billed at a rate of $1.00 per linc.
All manuscripts for The Great Lakes Entomologist should be sent to the Editor, Mark F. O'Brien, Insect
Division, Museum of Zoology, The University of Michigan, Ann Arbor, MI, 48109-1079, USA. Other
correspondence should be directed to the Executive Secretary (see inside front cover).

Vol. 24 No. 1 - Michigan Entomological Society

Transcription

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