The GREAT LAKES ENTOMOLOGIST · 2014. 12. 9. · THE GREAT LAKES ENTOMOLOGIST CORlXlD TRANSIENTS ON STREAM RIFFLES (HEMIPTERA: CORIXIDAE)' Ralph D. ~toaks.~ Joe K. eel,^ and Richard

The

GREAT LAKES ENTOMOLOGIST

Vol. 13, No. 2 Summer 1980

THE GREAT LAKES ENTOMOLOGIST

Volume 13

Published by the Michigan Entomological Society

No. 2 ISSN 0090-0222

TABLE OF CONTENTS

Corixid Transients on Stream Rimes (Hemiptera: Corixidae) R. D. Stoaks. J. K. Neel, and R. L. Post . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

An Annotated List of the Checkered Beetles (Coleoptera: Cleridae) of Michigan D. C. L. Gosling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

Size Reduction Southward in Michigan's Mustard White Butterfly, Pieris napi (Lepidoptera: Pieridae)

W. H. Wagner, Jr. and Michael K. Hansen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ?7

Simulation of the Effects of Stand Factors on Spruce Budworm (Lepidoptera: Tortricidae) Larval Redistribution

W. P. Kemp, J. P. Nyrop, and G. A. Simmons.. ............................. 81

Fleas of the Norway Rat in the Superior, Wisconsin, Harbor Area (Siphonaptera: Ceratophyllidae, Hystrichopsyllidae)

Darol L. Kaufrnan and Norman D. Radtke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

COVER ILLUSTRATION

Leptostylus transversus (Gyllenhal) (Coleoptera: Cerambycidae) on its host plant. poison ivy (Toxicodendron radicans). Photograph by N. M. Gosling

THE MICHIGAN ENTOMOLOGICAL SOCIETY

1979-80 OFFICERS

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M. C. Nielsen D. C. L. Gosling

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Copyright 1980, The Michigan Entomological Society


CORlXlD TRANSIENTS ON STREAM RIFFLES (HEMIPTERA: CORIXIDAE)'

Ralph D. ~ t o a k s . ~ Joe K. eel,^ and Richard L. post4

ABSTRACT

Five genera and 12 species of corixids were collected from riffle regions of the Forest River, North Dakota, from summer of 1970 to fall 1971. Their occurrence, ecology, succes- sion, and seasonal abundance in headwaters is discussed.

Corixids have been infrequently reported from lotic waters (Hilsenhoff 1972). Brooks and Kelton (1967) described the ecology, range, and distribution of corixids in the Prairie Prov- inces and presented generic keys applicable to the North Dakota species discussed below. Of the 41 species reviewed by Brooks and Kelton, none were reported from fast-water habitats or gravel bottom streams. However, 12 species of corixids in five genera were collected from headwaters of the Forest River in northeastern North Dakota during an investigation of riffle insects (Stoaks 1975). The river originates in Walsh and Nelson counties, courses eastward approximately 97 km, then discharges into the Red River of the North. Substrates were mainly sand and gravel with some slate, rocks, mud, and boulders. Headwater discharge declined before and after spring floods. The extreme increase in spring discharge (Table 1) resulted from melting ice. Because headwater reaches of the Forest River are typical riffle habitats (Brown and Shoemake 1964), corixids were unexpected. Nevertheless, corixids were found as facultative riffle inhabitants with greatest abundance and diversity in fall 1971.

Table I. Mean monthly and seasonal discharge rates at US Geological Survey gage on the Main Branch of the Forest River, North Dakota, 0.8 km below the confluence of the South Branch.

Month Discharge Ratea Month Discharge Ratea

June 0.56 December 0.24 July 0.44 January 0.20 August 0.20 February 0.25 September 0.21 March 0.45 October 0.40 April 12.88 November 0.32 May 1.07

June 0.61 July 2.00 August 0.40 September 0.30 Sovember 0.41

a V a l ~ 2 ~ in m3i5ec converted from CS Gml- Sun-e) 1190. 1971. 1972).

' ~ ~ ~ r o b ~ e d b the Director. Sonh Dakota Agricultural Experiment Station as Journal Article 1036. Article based on part of a dissertation b>- t k senior author and approved by the Entomology Department, Sorth Dakota Stare L-niversin .

?6711 Northwest h-b~e. Des Sfoines. IA 50311 3 ~ i o l ~ Department. Lnibersit) of North Dakota. Grand Forks. ND 58201 4~n tomolm Department. Zorth Dakota State University, Fargo, ND 58102.

THE GREAT LAKES ENTOMOLOGIST Vol. 13, No. 2

METHODS

Weekly to monthly diurnal collections were made with a Surber sampler at seven sampling stations of the Forest River from highway 18 to 32. Sampling was qualitative during summer and fall of 1970 and quantitative from winter 1970 to fall 1971 (Table 2). Four to 17 hauls were made for each collection. Detailed descriptions and maps of the collecting sites have been presented elsewhere (Stoaks 1975). Permanent slides of palae, strigilae, and hemelytrae of species identified in this study were deposited in the North Dakota Insect Survey collection.

DISCUSSION

Maximum corixid density was obtained from Surber samples taken in running water of 0.34 m3/sec velocity at a depth of 0.5 m, behind a submerged stump located nearly halfway across the Main Branch on 16 October 1971. Three hauls yielded 4314 specimens of the following species: Callicorixa audeni (87%), Sigara decoratella (6%). S. grossolineata (4%), Hesperocori-ra uulgaris (2.5961, H . michiganerzsis (1.2%). H. atopodonta (0.8%). and S. conocephala (0.2%). An analysis of quantitative data from winter of 1970 to fall 1971 indicates C . audeni was the fall dominant species at seven collecting stations at the headwaters of the Forest River inclusive of the North, Middle, South, and Main branches. C . audeni was replaced in summer by S. mathesoni and S. grossolineata (Table 2). H . vulgaris had greatest density in winter and spring. Other corixids collected are listed in Table 2. Corixid presence in a riffle habitat was related to the following conditions: (I) reduced discharge at riffles, especially in late summer and fall when drought caused drying of floodplain pools and formation of pools and intermittent stream conditions in the channel of the Middle and South branches of the river; (2) the Whitman Dam on the Middle Branch stopped discharge during the driest months and caused pool formation when ground water recharge was minimal; (3) accumulation of twigs, leaves, and other vegetation in riffles which simulated pool-like conditions; (4) peak populations occurred most frequently in fall when migratory activity is greatest in corixids seeking overwintering sites; (5) tolerance to over-icing conditions because of continuous recharge from aquifers having warmed waters. As drought extended into fall, corixid increases in tributaries were minimal compared to downstream. The declin- ing discharge began after spring run-off and tributaries tended to go dry before reaching their

Table 2. Seasonal abundancea of Corixidae in headwaters of the Forest River, North Dakota - - - -

1970 1971 -

Winter Spring Summer Fall Total

Callicorixa audeni Hungerford Cenocorixa bi'da (Hungerford) Hesperocorixa michisanensis (Hungerford) H . atopodonta Hungerford H . vulgaris (Hungerford) Sigara bicoloripennis (Walley) S . conocephala (Hungerford) S . decoratella (Hungerford) S . grossolineata Hungerford S . solensis (Hungerford) S. mathesoni Hungerford Trichocorixa nais Kirkaldy and Torre-Bueno

amean/Surber haul

1980 THE GREAT LAKES ENTOMOLOGIST 63

related aquifers. Corixids found in drying pools of riffles were S. grossolineata and S . decoratella. The greatest numbers of corixids were found at the riffle farthest downstream under conditions that probably best simulated a true corixid habitat. This was the site with least gradient, smooth discharge across riffles, and a large stump in mid channel. Even during winter, springs from aquifers along the North, South, and main branches maintained flow and kept ice thin or melted. Overwintering corixids were: C . audeni, H. vulgaris. S . bicoloripennis, S . mazhesoni, and S . grossolineata. From recent drift analyses in a separate study (unpublished), Neel learned that corixids have limited movement down riffles during periods of continuous discharge, but sometimes appear on rime heads in large numbers. Corixids found in the Forest River were transients into riffles and occurred under unique conditions described herein.

LITERATURE CITED

Brown, H. P. and C. M. Shoemake. 1%4. Oklahoma riffle beetles (Coleoptera: Dryopoi- dea). IV. Ecology. Proc. Oklahoma Acad. Sci. 44:4446.

Brooks, A. R. and L. A. Kelton. 1967. Aquatic and semiaquatic Heteroptera of Alberta, Saskatchewan, and Manitoba (Hemiptera) Mem. Entomol. Soc. Canada 51:7:92.

Hilsenhoff, W. L., J. L. Longridge, R. P. Narf, K. J. Tennessen, and C. P. Walton. 1972. Aquatic insects of the Pine-Popple River, Wisconsin. Tech. Bull. 54, Dept. Natur. Re- sourc. Madison.

Hungerford, H. B. 1948. The Corixidae of the Western Hemisphere. Univ. Kansas Sci. Bull. 32: 1-727.

Stoaks, R. D. 1975. Seasonal and spatial distribution of riffle dwelling aquatic insects in the Forest River, North Dakota. Ph.D. dissert. North Dakota State Univ., Fargo.

U.S. Geological Survey. 1970. Water resources data for North Dakota. Part I . Surface water records. U.S. Govt. Print. Office. Wash., D.C.

. 1971. Water resources data for North Dakota. Part I. Surface water records. U.S. Govt. Print. Office. Wash., D.C.

. 1972. Water resources data for North Dakota. Part I. Surface water records. U.S. Govt. Print. Office. Wash., D.C.

1980 THE GREAT LAKES ENTOMOLOGIST

AN ANNOTATED LIST OF THE CHECKERED BEETLES (COLEOPTERA: CLERIDAE) OF MICHIGAN

D. C. L. Gosling1

The checkered beetles (Cleridae) have been included in local lists of Michigan Coleoptera by Andrews ( 1916, 192 1 , 1929), Hatch (1924), Hubbard and Schwarz (1878). LeConte (1850), and Wolcott (1909). The present list is the first to include records from throughout the state, and is based on the collections of the University of Michigan Museum of Zoology (UMMZ), those of the Department of Entomology, Michigan State University (MSUC), and the private collections of E. F. Giesbert, D. K. Young, and the author. This list records 35 species of Cleridae from the state, and I believe it is essentially complete, although future collecting in the southern counties may well produce new records.

The annotations for each species include the known distribution in Michigan, the range of reported collection dates for adults in the state, a brief summary of recorded prey rela?ion- ships, and comments on collecting methods. Distribution data are shown in Figures 2-29, except for a few species known from only one or two counties. Open symbols represent published records not duplicated in the collections I have examined. The systematic order follows Barr (1975).

All of the species recorded in this list as occurring in Livingston County have been collected at the Edwin S. George Reserve, a research facility of the University of Michigan Museum of Zoology.

The larvae and adults of most species of Cleridae are important predators of bark beetles and wood-boring insects attacking both hardwoods and conifers. A few species are also recorded as preying on various gall insects. Three species of Michigan tlerids, Leconrella cancellara, Placopterus rhoracicus, and Trichodes nuralli, prey on Hymenoptera, some of which are beneficial insects. The three species of Necrobia combine feeding on camon with predation on other saprophagous insects. Most clerids, however. are considered beneficial natural enemies of many of our important forest pests.

Adult clerids may be collected by careful inspection of dead trees and logs, where they are found both beneath and on the surface of the bark. A headlight greatly facilitates collecting nocturnal species. Beating dead branches is another useful collecting method, and a few species may be taken by sweeping foliage and flowers. Some clerids have been collected in Malaise traps, and several species are attracted to lights. Specimens are often reared from logs and branches infested by wood-borers and bark beetles. Caution should be taken in infemng prey relationships for reared specimens, however, as several potential prey species are often present.

Knull (195 1) presented more complete biological information, keys, and excellent illustrations of all the species recorded in this list except Enoclerus leconrei. This useful and inexpensive bulletin has been reprinted and is available from the Ohio Biological Survey, 484 West 12th Avenue, Columbus, OH 43210. A description of E. leconrei is included in the annotation for that species.

Family CLERIDAE Subfamily THANEROCLERINAE

Genus ZENODOSUS Wolcott

sanguineus (Say) 1835. (Fig. 2). April to early September. Prey: small wood-borers infesting hardwoods. I have collected sanguineus on oak logs.

'~chool of Natural Resources, University of Michigan, Ann Arbor, MI 48109

66 THE GREAT LAKES ENTOMOLOGIST Vol. 13, No. 2

WISCONSIN

Fig. 1 . The counties of the State of Michigan

Subfamily CLERJNAE Genus MONOPHYLLA Spinola

terminata (Say) 1835. (Fig. 3). Mid-April to mid-June. Prey: Bostrichidae, Buprestidae, Cerambycidae, Lyctidae, and Scolytidae infesting hardwoods. I have reared terminata from walnut logs.


Genus LECONTELLA Wolcott and Chapin

cancellata (LeConte) 1854. Mid-July to mid-August. A rarely collected species known in Michigan only from Kalamazoo and St. Joseph counties. It has been recorded from the nests of several species of Hymenoptera (Foster and Barr 1972, Rau 1944).

Genus CYMATODERA Gray

bicolor (Say) 1825. (Fig. 4). June to mid-August. Prey: Anobiidae, Cerambycidae, and Scolytidae infesting juniper and various hardwoods. I have reared bicolor from juniper branches and hickory logs, and collected adults at UV lights in oak-hickory woodlands.

inornata (Say) 1835. (Fig. 5). Mid-June to mid-August. Prey: Anobiidae, Bostrichidae, Buprestidae, and Eucnemidae infesting hardwoods.

undulata (Say) 1825. (Fig. 6). Mid-July to late September. Prey: Cerambycidae, Cynipidae, and Eucnemidae infesting hardwoods.

Genus PRIOCERA Kirby

castanea (Newman) 1838. (Fig. 7). Late July to mid-August. Prey: Lymexylonidae infesting oak and Scolytidae infesting pine. I have collected this handsome species at UV lights in an oak-hickory woodland.

Genus PLACOFTERUS Wolcott

thoracicus (Olivier) 1795. (Fig. 8). Late May to late July. Prey: Cerambycidae, Eucnemidae, and Scolytidae infesting dead branches of hardwoods. I have reared this species from oak twigs infested by Eluphidionoides parallelus (Newman) (Cerambycidae), and beaten it from dead sumac branches. It is frequently attracted to UV lights in oak-hickory woodlands. Foster and Barr (1972) reported thoracicus from the nests of two species of wasps.

Genus THANASIMUS Latreille

dubius (Fabricius) 1776. (Fig. 9). Mid-May to early September. Prey: Scolytidae infesting conifers. I have collected dubius on pine logs and in traps baited with turpentine. The biology of this species is of current interest owing to its potential as a predator of the southern pine beetle, Dendroctonus frontalis Zimmermann (Scolytidae) (Dix and Franklin

68 THE GREAT LAKES ENTOMOLOGIST Vol. 13. No. 2

1977, Dixon and Payne 1979, Mignot and Anderson 1969, Reeve et al. 1980, Richerson et al. 1980, Thatcher and Pickard 1966, Turnbow and Franklin 1979, Turnbow et al. 1978, Vite' and Williamson 1970).

trifasciatus (Say) 1825. Early to mid-July. Prey: Cerambycidae and Scolytidae infesting conifers. I collected trifasciatus from a red pine log near Conway Lake in Marquette County. The only other Michigan specimen I have seen was collected at Deward in Crawford County (MSUC).

undatulus (Say) 1835. (Fig. 10). Mid-June through July. Prey: presumably Scolytidae infesting conifers. I have collected this species from pine logs and in beach drift along Lake Superior.

I980 THE GREAT LAKES ENTOMOLOGIST 69

Genus ENOCLERUS Gahan

[analis (LeConte) 1849. See Enocler~is rosmarus.] [ichneumoneus (Fabricius) 1776. See Enoclerus murrkowskii.] lecontei Wolcott 1910. Prey: Scolytidae infesting conifers. Wickham and Wolcott (1912)

recorded leconrei from Michigan but I have seen no specimens from the state in the collections I have examined. It can be distinguished easily from other Michigan species of Enoclerus by the uniform black coloration and absence of prominent white or light- colored fasciae on the elytra. The only elytral markings are three quite variable bands of sparse, gray pubescence. The basal fascia does not cover the humeri but may extend posteriorly to merge with a narrow, usually incomplete median fascia. The anterior margin of the apical fascia is transversely truncate from the suture to the middle of each elytron and then angulate posteriorly. Information on the biology of lecontei has been reported by Berryman (1966), Person (1940), and Rice (1971).

muttkowskii (Wolcott) 1909. (Fig. 11). Late June through July. Prey: Scolytidae infesting pine. I believe Wickham's (1895) record for ichneumoneus from the Lake Superior shore of Wisconsin probably refers to muttkowskii.

nigrifrons (Say) 1823. (Fig. 12). Early June to mid-August. Prey: Cerambycidae and Scoly- tidae infesting conifers and hardwoods. I have beaten this species from dead branches of sumac, larch, red pine, and jack pine, and reared it from larch branches.

nigripes (Say) 1823. (Fig. 13). April through July. Prey: Curculionidae and Scolytidae infesting conifers and hardwoods. King and Fox (1970) found that the color of the abdomen may not be a reliable character to use in distinguishing this species from nigrifrons.

quadrisignatus (Say) 1835. Late July. Prey: Scolytidae infesting elm and hickory. The only specimen from Michigan I have seen was collected in Monroe County by James Truchan from a Malaise trap (MSUC).

rosmarus (Say) 1823. (Fig. 14). Early May through July. I have beaten this species from dead sumac branches. Knull(1951) described it as "abundant on flowers and weeds." All of the specimens of rosmarus in the Andrews collection (MSUC, UMMZ) were incorrectly identified by him as analis. I believe that analis does not occur in Michigan, and presume that Andrews' (1916) record for analis from the Charity Islands in Saginaw Bay refers to rosmarus. However, I have not been able to locate his specimen to verify this.


Genus TRICHODES Herbst

nutalli (Kirby) 1818. (Fig. 15). Mid-June to early September. The larvae of nutalli are nest predators of bees and wasps. Adults are collected from various flowers where they feed on pollen. It is a common clerid in the northern part of Michigan, and particularly abundant in the Northern Lower Peninsula.

Subfamily PHYLLOBAENINAE Genus PHYLLOBAENUS Dejean

humerali (Say) 1823. (Fig. 16). Late May to early September. This species is generally collected by sweeping or beating foliage. The Marquette County record by Hubbard and Schwarz (1878) possibly refers to Phyllobaenus lecontei.


lecontei Wolcott 1912. (Fig. 17). Mid-June to mid-August. I have collected leconreifrom the flowers of Daucus carora L.

pallipennis (Say) 1825. (Fig. 18). July through September. A rather common species collected by beating dead branches, especially sumac and oak. It is also taken in Malaise traps and occasionally at UV lights. I. J. Cantrall and I have rearedpallipennis from oak logs.

unifasciatus (Say) 1825. (Fig. 19). July to mid-August. Prey: Buprestidae, Cerarnbycidae, Curculionidae, and Scolytidae infesting hardwoods. I have collected unifasciarus by beating dead sumac branches and reared it from walnut logs.

verticalis (Say) 1835. (Fig. 20). Late May to mid-July. Prey: Buprestidae, Cerambycidae, and Cynipidae infesting hardwoods.


~ H Y L L ~ B A E N U S P A L L I P E N N I S

1 a PHYLLDBAENUS V E R T I C A L I S

Genus ISOHYDNOCERA Chapin

curtipennis (Newrnan) 1840. (Fig. 21). Late May through August. Prey: Gelechiidae infesting Solidago and Tenthredinidae infesting willow.

tabida (LeConte) 1849. (Fig. 22). Early June through mid-July. Prey: Mordellidae infesting herbaceous plants.

Genus WOLCOTTIA Chapin

pedalis (LeConte) 1866. (Fig. 23). Late June to early July. Knull(1951) described this species as "abundant in old fields on weeds" but it has not been cbllected often in Michigan.

I980 THE GREAT LAKES ENTOMOLOGIST 73

Subfamily KORYNETINAE Genus PHLOGISTOSTERNUS Wolcott

dislocatus (Say) 1825. (Fig. 24). Early May through July. Prey: Buprestidae, Cerambycidae, Colydiidae, and Scolytidae infesting branches of hardwoods and conifers. I have collected dislocatus by beating dead branches of oak, sumac, and pine, and reared it from small larch logs.

Genus NEICHNEA Wolcott and Chapin

laticornis (Say) 1835. (Fig. 25). Late June to late July. Prey: Scolytidae infesting hardwoods and conifers. I have beaten laticornis from dead oak branches.


Genus NEORTHOPLEURA Barr

thoracica (Say) 1823. (= Orthopleura damicornis of Knull [Barr 19761). (Fig. 26). June to mid-August. Prey: Buprestidae, Cerambycidae, and Curculionidae infesting hardwoods. This species is often attracted to UV lights in oak-hickory woodlands.

Genus CHARIESSA Perty

pilosa (Forster) 1771. (Fig. 27). Late May to early August. Prey: Scolytidae infesting hardwoods and conifers. I have reared pilosa from hickory logs and collected it at UV lights in oak-hickory woodlands.

Genus CREGYA LeConte

oculata (Say) 1835. Early July through early August. Prey: Cerambycidae and Scolytidae infesting hardwoods and conifers. This species has been collected in Michigan only in Oakland and St. Joseph counties. I have taken it by beating dead sumac branches and those of other hardwoods, and at UV lights in oak-hickory woodlands.

Genus PELONIUM Spinola

leucophaeum (Klug) 1842. (= Corinthiscus leucophaeus of Knull [Barr 19751). Early July. The only Michigan specimen I have seen was collected in Wayne County by A. W. Andrews (MSUC).

Genus NECROBIA Olivier

Three cosmopolitan species of Necrobia have been collected in Michigan. They are usually found on the carcasses of dead animals where they feed both on the decaying canion and on other saprophagous insects such as Dermestidae and Sarcophagidae (El-Mallakh 1978). These clerids also are found as pests feeding on stored animal and plant products.

ruticollis (Fabricius) 1775. April. This species has been collected in Michigan only in Wayne and Washtenaw counties.

rufipes (DeGeer) 1775. (Fig. 28). June to late August. violacea (Linnaeus) 1758. (Fig. 29). Early April to mid-September.


ACKNOWLEDGMENTS

I would like to thank Dr. John A. Witter, School of Natural Resources, and Dr. Irving J. Cantrall, Museum of Zoology, University of Michigan, for their helpful review and comments on the manuscript; and Edmund F. Giesbert and Daniel K. Young for allowing me to include records from their private collections.

LITERATURE CITED

Andrews, A. W. 1916. Results of the Mershon Expedition to the Charity Islands, Lake Huron: Coleoptera. p. 65-108 in: Miscellaneous papers on the zoology of Michigan. Michigan Geol. and Biol. Surv. Publ. 20.

. 1921. The Coleoptera of the Shiras Expedition to Whitefish Point, Chippewa County, Michigan. Pap. Michigan Acad. Sci., Arts, and Letters. 1:293-390.

. 1929. List of some of the insects found at Huron Mountain. p. 116152 in: B. H. Christy (ed.). The book of Huron Mountain. Huron Mountain Club, Chicago.

Barr, W. F. 1975. Cleridae, the checkered beetles. p. 66.146.18 in: Checklist of the beetles of Canada, United States, Mexico, Central America and the West Indies: Volume I, Part 5, The darkling beetles, ladybird beetles, and related groups. Biol. Res. Inst. Amer., Latham, New York.

. 1976. Taxonomy of the new clerid genus Neorthopleura (Coleoptera). Melan- deria. 24: 1-14.

Benyman, A. A. 1%6. Studies on the behavior and development of Enoclerus lecontei (Wolcott), a predator of the western pine beetle. Canadian Entomol. 98519-526.

Dix, M. E. and R. T. Franklin. 1977. Diel activity of Thanasimus dubius, a southern pine beetle predator. J . Georgia Entomol. Soc. 12:71-75.

Dixon, W. N. and T. L. Payne. 1979. Aggregation of Thanasimus dubius on trees under mass-attack by the southern pine beetle. Environ. Entomol. 8:178-181.

El-Mallakh, R. 1978. A food source for Necrobia violacea (Coleoptera: Cleridae). Entomol. News. 89: 178.

Foster, D. E. and W. F . Barr. 1972. Notes on the distribution and bionomics of some North American Cleridae. J. Kansas. Entomol. Soc. 45:122-125.


Hatch, M. H. 1924. A list of Coleoptera from Charlevoix County, Michigan. Pap. Michigan Acad. Sci., Arts, and Letters. 4543-586.

Hubbard, H. G. and E. A. Schwarz. 1878. The Coleoptera of Michigan. Roc. Amer. Phil. SOC. 17593469.

King, W. E. and R. C. Fox. 1970. On the taxonomy of clerid species in South Carolina (Coleoptera: Cleridae). Roc. Entomol. Soc. Washington. 72:133.

Knull, J. N. 1951. The checkered beetles of Ohio (Coleoptera: Cleridae). Ohio Biol. SUN. Bull. 42. 7:267-350.

LeConte, J. L. 1850. General remarks upon the Coleoptera of Lake Superior. p. 201-242 in: L. Agassiz. Lake Superior: its physical character, vegetation, and animals, compared with those of other and similar regions. Gould, Kendall, and Lincoln, Boston.

Mignot, E. C. and R. F . Anderson. 1%9. Bionomics of the bark beetle predator Thanasimus dubius Fab. (Coleoptera: Cleridae). Entomol. News. 80:305-310.

Person, H. L. 1940. The clerid Thanasimus lecontei (Wolc.) as a factor in the control of the western pine beetle. J. Forestry. 38:3%396.

Rau, P. 1944. A note on Leconrella cancellara Lec. (Coleoptera: Cleridae) in cells of the mud-daubing wasp. Entomol. News. 13:61.

Reeve, R. J., J . E. Coster, and P. C. Johnson. 1980. Spatial distribution of flying southern pine beetles (Coleoptera: Scolytidae) and the predator Thanasimus dubius (Coleoptera: Cleridae). Environ. Entomol. 9: 113-1 18.

Rice, R. E. 1971. Flight characteristics of Enoclerus lecontei, Ternnochila virescens, and Tomicobia ~ibialis in central California. Pan-Pacific Entomol. 47:l-S.

Richerson, J. V., F. A. McCarty, and T. L. Payne. 1980. Disruption of southern pine beetle infestations with Frontalure. Environ. Entomol. 9:%93.

Thatcher, R. C. and L. S. Pickard. 1966. The clerid beetle, Thanasimus dubius, as a predator of the southern pine beetle. J. Econ. Entomol. 59:955-957.

Turnbow, R. H. and R. T. Franklin. 1979. Hyalomyodes rriangulifera (Diptera: Tachinidae): A parasite of the southern pine beetle predator Thanasim~is d~ibius (Coleoptera: Cleridae). J . Georgia Entomol. Soc. 14: 174-176.

Turnbow, R. H., R. T. Franklin, and W. P. Nagel. 1978. Prey consumption and longevity of adult Thanasimus dubius. Environ. Entomol. 7:695497.

Vit6, J. P. and D. L. Williamson. 1970. Thanasimus dltbius: Prey perceptions. J. Insect Physiol. 16:233-239.

Wickham, H. F. 1895. A list of Coleoptera from the southern shore of Lake Superior. Roc. Davenport Acad. Nat. Sci. 6: 125-169.

Wickham, H. F. and A. B. Wolcott. 1912. Notes on Cleridae from North and Central America. Bull. Natur. Hist. State Univ. Iowa. 6:4947.

Wolcott, A. B. 1909. Supplementary list o f ~ s l e koyale beetles. p. 204215 in: C. C. Adams. An ecological survey of Isle Royale, Lake Superior. Michigan Geol. SUN. 1908.


SIZE REDUCTION SOUTHWARD IN MICHIGAN'S MUSTARD WHITE BUTTERFLY, PIERIS NAP1 (LEPIDOP'TERA: PIERIDAE)

W.H.Wagner, Jr. and Michael K. ~ a n s e n l

The mustard white, Pieris napi (Linnaeus) is generally considered to be a circumboreal species or species-complex. Much disagreement exists regarding the classification in this group, the members of which are extremely diverse, especially in Eurasia. There is evidently "a continuous range of differentiation, from local populations, through subspecies, to species, which nomenclature cannot fully reflect" (Bowden 1972). In North America the major controversy has been whether the so-called West Virginia white, P . virginiensis Edwards, should be separated as a distinct species. Most writers today agree that it should. Of the mustard white in the Great Lakes area, the local subspecies is genelally treated as P . napi oleracea Hams (for summary ofthe taxonomicproblems, cf. Howe [1975]). This taxonoccurs extensively in Canada, and down into Minnesota, Wisconsin, and Michigan, where it be- comes progressively less common southward. It is two-brooded, the members of the first brood commonly flying together with the univoltine virginiensis where their ranges overlap (Voss and Wagner 1956). Where they occur together, the two taxa are easily recognized and separated, but in the southern mountains, where napi is absent, the Appalachian form of virginiensis resembles the summer form of napi (Wagner 1978).

A familiar dogma in North American lepidopterology states that napi has been largely or altogether extirpated over much of its range by the cabbage white, P . rapae (Linnaeus) after the introduction of the latter around the middle of the 19th century. Whether or not this is really true needs further analysis, since many northern species of animals shifted their ranges farther .north, not because of competing sister species, but as a result of many decades of land-clearing and changing climatic conditions. If napi was reduced in numbers in the upper Great Lakes area it was probably due to this, and not the invasion of rapae. Possibly this has to do with the interesting size relationships to be described below. P . napi, in Michigan, just as virginiensis, appears to be a species of natural habitats, while the naturalized rapae is more a species of farmlands. However, in northern Michigan, as well as Wisconsin and Minnesota, all three species may be found flying together in mixed areas of farmland and native forest. In northern Michigan, as elsewhere in the upper Great Lakes, napi occurs in a variety of woodland and swamp habitats, including jack pine uplands. In southern Michigan, most of the colonies of napi, many of which are extensive and contain very large numbers of individuals, are found in cool swampy and marshy areas, often at the edges of bogs and fens.

In our studies of Michigan's Pierinae, we noticed what appeared to be an anomalous pattern of size variation which we report here. Although the usual trend in most groups of macrolepi- doptera is for clines of increasing size from north to south, the situation appears to be reversed in Michigan's napi. The largest individuals occur in the Upper Peninsula, some of them with forewings 25-26 mm long, and the smallest in the Southern Lower Peninsula, some with forewings down to 18-19 mm long. The smallest individuals seem to be concentrated in southern Michigan. These run about two-thirds the size of the largest Upper Peninsula individuals. In the vicinity of Ann Arbor we have found that very small individuals of only 19 or 20 mm forewing length are easily obtained in both the spring and summer broods, compris- ing up to approximately one-third and one-fifth of the populations respectively.

MATERIALS, METHODS, AND RESULTS

The material used for this study came from the collections of MogensC. Nielsen, Michigan State University, and W. H. Wagner, Jr. The length of the forewing, from the base of the

l~ iv i s ion of Biological Sciences, The University of Michigan, Ann Arbor, MI 48109


costal vein to the apex of the wing, was measured on a total of 112 specimens from25 Michigan counties. The measurements were then separated into groups on the basis of geographical location, either northern (north of Clare County-14 counties) or southern (south of Clare County-1 1 counties), and season of collection, whether spring or summer. The sexes were not separated in the data set, as there was no evidence of significant sexual dimorphism in size. As the number of specimens in each of the four groups was not equal, we chose at random 21 specimens from each group for statistical analysis in order to test the significance of the differences in size between north-south and spring-summer forms (Table 1). An analysis of variance (ANOVA) was also used (Table 2).

Table 1 gives the mean and standard deviation in forewing measurements for the four groups. On the average, the northern forms are larger than the southern forms and the summer forms are larger than the spring forms. The size difference between the northern and southern specimens is more pronounced in the spring than in the summer forms, the former averaging 2. IS mm longer, or 4.3 mm overall, in the north than in the south, an increase of 10.4%. For summer forms, the difference is less distinct, the northern specimens averaging 1.76 mm or 8% longer. As for the seasonal forms, compared inter se, the summer one is larger, with average difference in size in the north 0.81 mm or 1.6 mm overall, a 3.6% increase over the spring form, and in the south 1.2 mm or 2.4 mm overall, a 5.8% increase. The results of the statistical analysis thus show that the effects of season and geography on forewing lengths are both highly significant (Table 2).

DISCUSSION

The generally recognized north-south trend for increase in size in Great Lakes butterflies seems to be associated mainly with non-migratory species. Members of such butterfly genera as Papilio, Pieris, Cercyonis, Euptychia, Lethe, Boloria, Speyeria, Erynnis, Hesperia, and Poanes, for example, tend to show southward size increases. Similar observations may be made in certain moths, such as the saturniids Eacles, Antheraea, and Hyalophora. The reverse seems to be rare. Collins (1973) in studies of Hyalophora columbia (Smith) mentioned, that "Surprisingly, the moths from northern locales were larger on the whole in both average and maximum size than specimens taken farther south." He was referring to collec-

Table 1. Mean forewing lengths ( 5 SD) of spring and summer forms of P. napi in northern and southern Michigan.

Northern Michigan Southern Michigan

Spring Summer

Table 2. ANOVA for forewing lengths in P. napi.

Source SS d f MS F value

Locale 80.0476 1 80.0476 39.66%a Season 21.00 1 21.00 10.4071b Interaction .7619 1 ,7619 ,3776 Error 16 1.4286 80 2.0179 Total 263.2381 83

aEffect significant at the 0.01% level b~ffect significant at the 1% level.

THE GREAT LAKES ENTOMOLOGIST 79

Fig. 1 . Pieris napi oleracea, from Michigan, showing range of forewing length, 26 mm to 18 mm. Left column. top to bottom, springfo-top two, MackinacCo., bottom two, Washtenaw Co. Rightcolumn, top to bottom, summer form-top two, Cheboygan Co., bottom two, Jackson Co.

tions made several hundred miles north of Winnipeg compared to collections made east of - -

Winnipeg. It is, of course, not surprising that the spring broods of P. napi ate made up of butterflies

which average smaller than those of the summer brood. This is a general rule for bi- or multivoltine but tedies throughout temperate North America and elsewhere. Among other Michigan Pierinae, this seasonal trend is shown well in both the native checkered white, P. protodice (Boisduval and LeConte) and the naturalized cabbage white, P. rapae. Among univoltine Pierinae. we have found that the dunes form of the marbled white, Euchloe olympia (Edwards), runs smaller in Bemen County (also in Larnpton County, Ontario) than does the inland form of savannahs and prairies which flies at the same time (late April, May). The


smaller size of the dunes form of E. olympia may be a consequence of unusual climatic and edaphic conditions in the dunes: exposure, winds, permeable sands, and lowered growth potential in food plants (Wagner 1977).

In the present situation regarding the north-south decrease in size in Pieris napi, on the contrary, an explanation is not easy to postulate. For many reasons, including warmer temperatures, longer growth seasons, and richer flora, the normal expectation would be increase in size from small in the north to large in the south, the trend familiar in most other size-clines in Great Lakes butterflies. However, the same factors may act negatively in the case of napi. The progressively warmer climate toward the south end of the range may actually impede growth and development of this basically northern species. Other possible factors to consider include the nature of the food plants in north versus south; perhaps the larval food in southern Michigan involves different species and this causes a size difference. Also, historical factors related to the last glaciation may have played a role through possible isolation. For example, a separated population in the vicinity of glacial Lake Maumee may have drifted toward dwarfism and the effects of this may still be observable, even though the typical northern form has once again merged with the isolated one.

ACKNOWLEDGMENTS

We wish to thank Mogens C. Nielsen for permitting us to study specimens in his private collection, David J. Bay for making the photograph, and Roland L. Fischer, Curator of the Entomological Collections at Michigan State University for kindly making the materials of Pieris available.

LITERATURE CITED

Bowden, S. R. 1972. Pieris napi L. (Pieridae) and the superspecies concept. J. Lepid. Soc. 26: 17@-173.

Collins, Michael M. 1973. Notes on the taxonomic status of Hyalophora columbia (Saturnii- dae). J . Lepid. Soc. 27225-235.

Howe, William H. (ed.) 1975. The butterflies of North America. Doubleday and Co., Garden City, N.Y.

Voss, Edward G. and Warren H. Wagner, Jr. 1956. Notes on Pieris virginiensis and Erora laeta-two butteflies heretofore unreported from Michigan. Lepid. News 10: 18-24.

Wagner, Warren H., Jr. 1977. A distinctive dune form of the marbled white butterfly, Euchloe olympia (Lepidoptera: Pieridae) in the Great Lakes Area. Great Lakes Entomol. 10: 107- 112.

. 1978. The northern Great Lakes white, Pieris virginiensis (Lepidoptera: Pieridae) in comparison with its southern counterpart. Great Lakes Entomol. 1153-57.

THE GREAT LAKES ENTOMOLOGIST 81

SIMULATION OF THE EFFECTS OF STAND FACTORS ON SPRUCE BUDWORM (LEPIDOPTERA: TORTRICIDAE)

LARVAL REDISTRIBUTION1

W. P. Kemp, J. P. Nyrop and G. A. ~ i m m o n s ~

ABSTRACT

A model was developed to simulate the life history of instar-I through instar-I1 spruce budworm, Choristoneura fumiferana (Clemens). Particular emphasis was given to the redistribution phases during spring and fall and the factors affecting budworm survival. An- alyses included the effects of aggregated stand types, temperature, cloud cover, open areas, and the species distribution, on redistribution mortality. Of those stand factors examined, open area and non-host percentages were the most important in regulating redistribution survival.

The eastern spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortri- cidae), is native to North America and is distributed throughout the northern boreal forest from Alberta to Maine. The preferred hosts are balsam fir (Abies balsamea [L.] Mill.), white spruce (Picea glauca [Muench] Voss) red spruce (P. rubens Sarg.), and to a lesser extent black spruce (P. mariana [Mill.] B.S.P.) (Greenbank 1963). All species are harvested for pulpwood and represent an economic base in Quebec, New Brunswick, and Maine.

Outbreaks of the budworm occur in 3 W 0 year intervals. The outbreaks themselves may extend only 8-10 years, but the effects are longer lasting (Blais 1965; Fye and Thomas 1963; Ghent et a1 1957; Ghent 1958, 1963; Hatcher 1961, 1963, 1%4; Heimberger 1945; McLintock 1955; Williams 1966). In the past, large scale spraying programs reduced budworm populations. However, concerns have been voiced about the effectiveness and environmental impact of the strategies (Blais 1974). Studies on spruce budworm dynamics and forest character were suggested and in some cases attempted. These new studies were concerned with the interactions of budworm and the forest, emphasizing possible unique means of control.

Among studies initiated were those aimed at manipulating stand structure and density to regulate sub-outbreak populations (Simmons et al. 1975; Kemp 1978; Kemp and Simmons 1978, 1979). These studies suggested that stands indeed influence budworm mortality factors. Further field studies, however, will require upwards of 25 to 50 years to apply these findings. In order to better design long-term field studies, a simulation model was used to help identify promising ways in which stands can be manipulated for increased mortality during larval redistribution periods. Two periods were examined: (1) the period following egg hatch when larvae disperse but do not feed, and (2) the period following emergence from overwintering hibernaculae when larvae redistribute then establish feeding sites.

Few procedures have been developed to model larval redistribution in a heterogenous environment. Watt (1968) developed and Kitching (1971) used a system where inputs and outputs for each discrete habitat unit were recorded and summed for each time interval in the dispersal period. For this case, redistribution was modeled within an assemblage of trees rather than as a process influenced by individual trees. With this method, the effects of stand composition and spatial pattern on redistribution were investigated with greater ease. Using the theory of the nearest neighbor measure of spatial relations, a model was developed, incorporating past and present information, to define effects of stand composition, spacing, and weather on the losses of spruce budworm during the fall and spring redistributions.

I ~ i c h i ~ a n Agricultural Experiment Station Journal Article No. 8635. 2~e~ar tmenl of Enlomology, Michigan State University, East Lansing, MI 48824.

THE GREAT LAKES ENTOMOLOGIST Vol. 13. No. 2

MODEL DESCRIPTION-GENERAL

Diagrams of hypothetical stands were constructed by assigning a numeric value to each block on a grid designating either a host or a non-host tree. For each tree in the grid pattern, the probab!lity of encountering a host or non-host as a nearest neighbor in a hexagonal arrangement was calculated (Clark and Evans 1954). Averaging these calculations for the stand gave four conditional probabilities: the probability of encountering a host or non-host tree given that the starting point is a host tree and the same probabilities given that the starting point is a non-host tree.

A generalized redistribution model was formulated by first assuming that the host species (black, white, and red spruce, and balsam fir) were randomly dispersed throughout the areas designated as host. Open areas were distributed randomly throughout the stand. In addition, it was assumed that the probability of a larva hitting an open area was proportional to available open area. Finally, it was assumed that the probabilities of interception by a host o r non-host, calculated by evaluating the nearest neighbors. were the same for all distances travelled by the dispersing larvae (Fig. 1).

Since most redistribution occurs from directionless updrafts, the process was simulated in random directions. A proportion of larvae were assigned to hosts, non-hosts, and open areas as a function of the probability of intercepting each of these after the larvae left a source. Mortality occurred when larvae impacted open areas. Twenty percent of the larvae inter- cepted by non-hosts were also assumed to eventually reach the forest floor and die. Prior to each dispersal, the larvae were lumped as a common variable. A more specific description of fall and spring redistribution follows.

Fig. I . Driving routine for the spruce budworm redistribution model.


MODEL DESCRIPTION-SPECIFIC

In the fall and spring, the number of larvae redistributing is a proportion of the total available to redistribute. A number of factors contribute to determining this proportion. Stand density and cloud cover influence both fall and spring redistribution. Larval dispersal occurs when the photopositive larvae (Wellington 1948) crawl to the tips of the branches, spin silk threads, and are carried by the wind (Henson i950). Two factors limiting redistribution are the number of larvae responding photopositively and turbulent wind conditions capable of launching larvae.

First-instar budworm emerge from the egg mass over a two week period in any one area (Mott 1%3). Cloud cover determines the actual number dispersing each day. In the model, the redistribution period was aggregated into one redistribution (Fig. 2). The effect of weather was simplified where the proportion of dispersing larvae was dependent on the number of (clear) dispersal days. This is a valid simplification of Shaw and Little (1973) who found virtually no redistribution occurring on overcast days; thus weather effect was ex- pressed as a percentage of favorable days during the redistribution period. Spring dispersal occurs repeatedly, depending on the length of time required to accumulate a specified number of degree days. In this case, the proportional effect of weather was divided by the number of dispersals occurring (Fig. 3).

The transfer of small larvae from tree to tree or from tree to open area depends on air currents breaking silk threads spun by the larvae. A greater proportionwould be expected in open stands which permit free air movement facilitating updrafts. Morris and Mott (1%3) found that approximately 50% of the larvae dispersed in dense stands and almost all dispersed in open stands. Since stand stocking in the model is held constant, stand density must

FALL REDISTRIBUTION (OCCURS ONCE IN MODEL)

PARASITISM

EGG HATCH

C L O COVER

WIND REDISTRWTION

TWIBULENCE

TEMPERATURE

v OVER-

WINTERING

Fig. 2. Fall redistribution subroutine illustrating major components and interactions.


SPRING REDISTRIBUTION

(OCCURS REPEATEDLEY IN MODEL)

HIBERNACULA WGl EMERGENCE PARASTISM

ACCUMULATION

FEEDING SITE

PREFERENCE

Fig. 3. Spring redistribution subroutine illustrating major components and interactions.

be defined in terms of the non-host trees that are leafless and contribute to the "openness" of the stand. The multiplier of larvae redistributing as influenced by stand density is defined as:

Y = 0.5 + (0.6 x NHOST) + (1.2 X OPEN) (1) NHOST and OPEN are the non-host and open components respectively. The proportion of larvae dispersing from the total available larvae, due to the effect of stand density and weather, is then:

DISLAR = TOTLAR x Y x WEATH (2)

DISLAR are the dispersing larvae, TOTLAR are the total larvae available, Y is the stand density factor with a range of 0.5 to 1.0, and WEATH is the proportion of cloud cover with a range 0.0 to 1.0. During spring redistribution WEATH is defined as: (WEATH)l(number of dispersals).

Parasitism by Trichogramma rninutum Riley, Apanteles fumiferanae Viereck and Glypta fumiferanae (Viereck) influences the number of dispersing larvae. T. minuturn is an egg parasite which reduces the total number of budworm available to disperse from the egg stage. G. fumiferanae and A. ficmiferanae, while not causing mortality in the first- or second-instars, alter the behavior of parasitized larvae and reduce the, number dispersing.


Parasitism by each species is dynamic and is influenced by stand composition and density. Kemp and Simmons (1978) found that T. rninuturn parasitism rates rose with increasing density of non-budworm host tree species. This was attributed to an increased availability of alternative hosts for T. rninuturn. In the model, G. furniferanae and A. furniferanae also increase with increased levels of open areas and non-hosts. Simmons et al. (1975) suggested that spruce budworm parasite densities are influenced by tree species density and composition. A more diverse forest with lower densities of spruce and fir provides a greater variety of alternate host insects. Openings in the canopy facilitate support of herbaceous plants on the forest floor providing nectar, an important food of adult parasites (Syme 1966, 1975; Price 1976; Leuis 1%7). In addition, these openings retain the parasites within the stand, reduce energy expenditures needed for successful host exploitation, and increase the chances of parasitism (Simmons et al. 1975). Using information presented by Kemp and Simmons (1978), T. rninuturn parasitism is given as:

PAR = 0.01 + (NHOST X 0.58) (3)

G. furnifrranae and A. furnifrranae parasitism are given as:

PAR = 0.1 + (0.3 x NHOST) + (0.55 x OPEN) (4) Ranges for this rate were obtained from Simmons et al. (1977).

G. furnifrranae and A. furniferanae alter the behavior of parasitized larvae by changing the phototactic response from positive to negative (Wellington 1948). Lewis (1960) found that 25% of the parasitized larvae did not disperse. TOTLAR used in equation 2 was redefined for spring dispersal as the unparasitized hudwom plus 75% of the parasitized budworm:

TOTLAR = BUDW + (PARBUD x 0.75) (5)

where BUDW are the normal and PARBUD are the parasitized budworm. PARBUD is determined:

PARBUD = TOTBUD x PAR (6)

and BUDW is:

BUDW = TOTBUD - PARBUD (7)

Where fall dispersal occurs only once in the model, spring dispersal takes place repeatedly (Figs. 2 and 3). Two factors interact to determine the number of larvae dispersing after the effects of cloud cover, stand density, and parasitism have been evaluated: the preferences for each host exhibited by larvae and the number of dispersals.

Shaw and Little (1973) reported that most budworm dispersal occurred during the warrn- est part of the day. Therefore, spring redistribution was modelled as the sum of dispersals taking place one each day over a length of time determined by temperature conditions. It was assumed dispersals would continue until larvae starved. Starvation results at 78 degree- days from the start of dispersal (Leonard, pers. comm.3). Thus, if the weather during the spring redistribution period was cool, more dispersals could occur than if it had been warm.

Jaynes and Speers (1949) measured the percentage of larvae reaching expanding buds in the spring. Those larvae not reaching the bud sites were assumed to have redistributed. The average of two reported values each for balsam fir, red spruce, and black spruce were 5696, 47%, and 1 I%, respectively. The ordering of these percentages corresponds to the reported budworm preferences for these species (Blais 1957). These figures and an interpolated value for the white spruce of 30% were used as an indication of budworm preference. These

3 ~ r . David E. Leonard, Dept. of Entomology, University of Maine, Orono, ME 04473


preferences change with increasing total degree days; when the starvation point is reached, none of the larvae disperse.

Third-instar budworm suffer greater mortality on black spruce due to late bud break (Blais 1957). In addition, larval development is slower (Blais 1957). Similar differences may occur on other host species, and thus the stimulus to redisperse may increase the chances of survival. At present, however, published data are not available so preference functions did not include this aspect. The preference function for all host species is given below:

Black spruce (BS) = .08 + (.00341 x TOTDD) Red spruce (RS) = .27 + (.00270 x TOTDD)

White spruce (WS) = .30 + (.00259 x TOTDD) Balsam fir (BF) = .56 + (.0016 x TOTDD)

The preference for the non-host trees is always 0. At the completion of redistribution larvae on non-hosts and 50% of those on black spruce perish.

Overwintering mortality as described by Miller (1958) was constant. While this is in no way indicative of a real-world situation, no data are currently available with which to model a dynamic winter mortality. Finally, degree-days were calculated each day using the method outlined by Baskerville and Emin (1969). Degree-days were used to determine the length of the spring redistribution period.

ANALYSIS

This model was utilized to (1) bring together as much information about the spruce budworm larval redistribution as possible and (2) aid in the development of future forest management methods to reduce spruce budworm outbreaks. Since the spruce budworm is most vulnerable during the fall and spring distribution periods, it follows that information of this nature could be incorporated in future management practices.

In this model, there were three periods of mortality: fall, winter, and spring. Winter mortality was constant throughout the model. This was defined by Miller (1958) as such. However, mortality in the fall and spring were both dynamic and based on a number of variables such as stand composition, stocking, and weather. Analyses, therefore, empha- sized the relationships involved in the fall and spring redistribution periods.

In order to effectively analyze the components of this model, three types of forests were established. These included non-host densities of approximately 9%, 4196, and 65%. Cor- responding host densities were 81%, 49%, and 25%. Open area was considered to be randomly distributed in both the host and non-host components of the stand. By altering different components of the model, we were able to examine (1) the effects of clumped versus random spacing of host and non-host species, (2) the effects of temperature, (3) the effects of cloud cover, (4) the effects of open area, and (5) the effects of host species composition on redistribution mortality.

RESULTS AND DISCUSSION

Figure 4A was the basis for all comparisons. Here spring redistribution mortality was greater than fall redistribution mortality. Further, spring redistribution losses showed different rates of mortality increase, where fall redistribution mortality showed a constant increase. These relationships remained true for all the variations tested. Figure 4A shows an increase in redistribution mortality in both the fall and the spring as nqn-hosts increase.

Stand spacing was the first component analyzed. By aggregating hosts and non-hosts, we found that fall and spring budworm redistribution mortalities were slightly reduced (Fig. 4B). Figure 4B illustrates the increased success of finding suitable hosts when budworm originate from suitable hosts.


Since temperature is considered an important factor in the spring dispersal process, the effects of a shorter dispersal time due to warm weather were examined. In Figure 4C all variables were held constant except daily temperatures. In this case, maximum-minimum temperatures from a warmer spring were used. Fewer dispersals result from warmer springs. Therefore, there is a decrease in the losses of budworms frorn.redistribution to unsuitable hosts in the spring.

Figures 4D and 4E, when compared with Figure 4A, indicate the effects ofcloudy weather on the redistribution losses. All components were held constant except that during the fall and spring redistribution 25% and 75% of the days were cloudy. Figure 4D shows a slight

100 . . . 100 . . . . . . .

0 20 40 m m o LO 40 60 W

A x wm ST 0 + NOW HOST

100 . . . . . . . 100 . . . . . . .

n o .

0 10 40 .O W 0 ' 20 40 10 a0

C x MH HOST D x MN HOST

. . . . . . I I I . . . I . . . . 0 1 0 4 0 6 0 1 0 0 LO 4 0 80 1 0

E Z Nul HOST F + WDI HOST

Fig. 4. Graph of the effects of non-host percentages on dispersal mortality of spring and fall budwom. (A) Weather = optimal, Open Area = 10%, Dispersal = 13, random spacing; (B) Host and non-host aggregated. (Fig. 4A superimposed in hatched lines B-J); (C) Dispersal time decreased by increasing temperatures; (D) Weather conlains 25% cloudy days; (E) Weather during dispersal contains 75% cloudy days; (F) Open area increased to 20%;


100 . I . . . . '

60 .

Z

0 20 +O 60 BO - G

0 10 1\7 w 80 % NOH HOST

H X W m w s ,

Fig. 4. continued. (G) Open area increased to 30%; (H) Open area reduced to (I) Species proportions were altered to increase black spruce; (J) Species proportions of the host component were altered to increase red spruce.

100

5 : ; I # 'O

reduction in mortality where Figure 4E shows a great reduction in spring mortality, when compared with optimal cloud conditions.

Figures 4 C 4 E indicate environmental effects that cannot be controlled. The effects can, in some cases, increase budworm mortality in forest stands. However, the components that can be manipulated are very important in that they are the only means we have for managing spruce budworm with silvicultural techniques.

As indicated earlier, Figure 4A shows that as non-hosts in a stand i n c ~ a s e , the percent mortality during the fall and spring redistribution periods increases. Not only do the non- hosts directly affect mortality from the interspersion of unsuitable feeding sites, but they also, in conjunction with open areas, influence parasitism rates which are important in fall mortality and spring redistribution. Figure 4A shows the effect of 10% open area. By increasing the open areas to 20% (Fig. 4F) and 30% (Fig. 4G), we found that there were rather large increases in mortality. This was true especially in the spring and at lower densities of non-host species. Figure 4H shows that by reducing the open area in the stand to W o , there was a marked reduction in fall and spring mortality. These figures indicate that open area is one of the most important components that can be manipulated.

Another forest component that can be manipulated is the species composition of the stand. In all previous figures this remained constant (BF = 0.60, RS = 1). 15, WS = 0. IS, BS = 0.10). However, in Figures 41 and 4J, the proportions of host species were changed to BF = 0.10, WS = 0.15, RS = 0.15, BS = 0.60, and BF = 0.15, WS = 0.15, RS = 0.60, BS = 0.10, respectively. Compared to Figure 4A, 41 indicates that by increasing the proportion of black spruce in the stand, spring mortality is greatly increased. This is owing to black spruce breaking bud late in the spring. Budworms dispersing to this host would tend to re-disperse.

. . .

// : : o Go n o ! : / , o 20 - w so eo !

I x IX*( HOST J % IPH HOST


As a result, survival of re-dispersing larvae is reduced. By contrast, Figure 45 shows that increasing the proportion of red spruce, though not the most preferred host, does not markedly increase mortality in the spring.

IMPLICATIONS IN FOREST MANAGEMENT

The results of this study indicate that there is a relationship between budworm survival during the spring redistribution period, the species composition, and the open area of the surrounding stand. This, combined with the observations of Turner (1952) and Batzer (1969, 1976) of reduced damage to fir as the proportion of hardwoods increases, lends credibility to the adoption of silvicultural techniques for management of spruce-fir in the northern boreal forest. First, by reducing the balsam fir in managed stands, we can more effectively manage against overwintering spruce budworm survival. The reduction of preferred budworm overwintering sites (balsam fir staminate flower bracts) in a spruce-fir stand would facilitate this population reduction. By moving away from a pre-climax or climax forest, we may also manage against budworm. Analyses indicate that in the spring, and to some extent the fall, redistribution losses can indeed be influenced by canopy openings and non-hosts. There- fore, management practices presently used in spruce-fir silvicultural systems, such as large clearcuttings and shelterwood methods, should be modified or replaced. Both clearcuttings and shelterwood techniques perpetuate even-aged stands of predominantly balsam fir (depending on site conditions). These stands are considered highly susceptible to budworm attack (Hatcher 1960). Selection cuttings, though more difficult, encourage uneven stand structure and at the same time allow for regulation of reproduction, canopy openings, and species composition. Using these methods, large susceptible aggregations of balsam fir could be discouraged while canopy openings and white spruce are encouraged. Non-host hardwoods in the maximum proportions economically allowed could be encouraged to further retard budworm outbreaks during the spruce-fir rotation. The diversification of the forest with canopy openings and non-host species would also serve to increase predators and parasites of budworm by increasing predator and parasite alternate host habitats (Simmons et al. 1975). This would be a very important, though indirect effect, of interspers- ing hardwoods and open areas in spruce-fir stands.

Probably not all stands are amenable to improved silvicultural practices. Large expanses of spruce-fir are owing in part to site conditions which are not tolerated by hardwoods. However, in some areas strip clearcuttings are used commercially and show potential in management of the forest against the budworm. In areas where accessibility is facilitated by road networks, patch clearcut harvesting is now being conducted. This method of harvesting also shows potential as a forest management practice that minimizes budworm impact. Small areas less than 10,000 acres that are intensively managed, as well as areas where there exists adequate varieties of species, both show excellent potential for management.

Improved silviculture likely cannot be viewed as a panacea, but certainly would contribute to forest management by providing additional methods that, when integrated with other control methods (i.e. chemical), would lead to improved spruce-firlspruce budworm management.

ACKNOWLEDGMENTS

We would like to thank Dr. Eric Goodman of the Department of Electrical Engineering and Systems Science and Dr. William Cooper of the Department of Zoology at Michigan State University for guidance and critical review. Funding of this project was made possible by a grant from the Maine Department of Conservation, Appropriation Number 1505.401 1.

LITERATURE CITED

Baskerville, G. L. and P. Emin. 1%9. Rapid estimation of heat accumulation from maximum and minimum temperatures. Ecology 50:51&517.


Batzer, H. 0. 1969. The forest character and vulnerability of balsam fir to spruce budworm in Minnesota. Forest Sci. 15:17-25.

. 1976. Silvicultural control techniques for the spruce budworm. p. 11CL116 in: Chansler, J. F . and W. H. Klein. Proceeding of a Symposium on the Spruce Budworm, November 11-14. 1974, Alexandria, Virginia. USDA Misc. Pub. 1327.

Blais, J . R. 1957. Some relationships of the spruce budworm Chorisroneura fumiferana to black spruce Picea ~nariana. Forest Chron. 33:364-372.

. 1%5. Spruce budworm outbreaks in the past three centuries in the Lauren- tide Park, Quebec. Forest Sci. 11: 13CL138.

. 1974. The policy of keeping trees alive via spray operations may hasten the recurrence of spruce budworm outbreaks. Forest Chron. 50:19-21.

Clark, P. J., and F. C. Evans. 1954. Distance to nearest neighbor as a measure of spatial relationships in populations. Ecology 35:445453.

Fye, R. E., and J. R. Thomas. 1963. Regeneration of balsam fir and spruce about fifteen years following release by spruce budwom attacks. Forest Chron. 39:385-397.

Ghent, A. W. 1958. Studies of regeneration in forest stands devastated by the spruce budworm. 11. Forest Sci. 4:135-146.

. 1963. Studies of regeneration in forest stands devastated by the spruce budworm. 111. Forest Sci. 9:295-301.

Ghent, A. W., D. A. Fraser and J. B. Thomas. 1957. Studies of regeneration in forest stands devastated by the spruce budworm. I. Forest Sci. 3:184-207.

Greenbank, D. 0. 1963. Host species and the spruce budworm. p. 219-223 in: R. F. Morris (ed.). The dvnamics of epidemic spruce budwom populations. Entoml. Soc. Canada - . ~ e h . 3 1: 1-j32.

Hatcher, R. J. 1960. Development of balsam fir following a clearcut in Quebec. Canadian Dept. North. Affairs and Natur. Resourc., Forest Resources Div., Tech. Note 87.

. 1961. Partial cutting balsam fir stands on the Epaule River watershed, Que- bec. Canadian Dept. Forest., Forest Res. Branch Tech. Note 105.

. 1%3. Effects of birch dieback and spruce budworm on forest development, Forest Section L. 6, Quebec. Canadian Dept. Forest., Forest Res. Branch, Pub. 1014.

. 1%4. Spruce budworm damage to balsam fir in immature stands, Quebec. Forest Chron. 40:372-383.

Heimberger, C. C. 1945. Comment on the budworm outbreak in Ontario and Quebec. Forest Chron. 21:114-126.

Henson, W. R. 1950. The means of dispersal of the spruce budworm. Ph.D. thesis Yale Univ., New Haven, Conn.

Jaynes, J. A., and C. F. Speers. 1949. Biological and ecological. studies of the spruce budwom. J. Econ. Entomol. 42:22 1-225.

Kemp, W. P. 1978. Stand parameters affecting mortality factors of spruce budwom eggs and dispersing larvae. M. S. thesis, Michigan State Univ.

Kemp, W. P. and G. A. Simmons. 1978. The influence of stand factors ,on parasitism of spruce budworm eggs by Trichogramma minutum. Environ. Entomol. 7:68%588.

. 1979. The influence of stand factors on survival of early instar spruce budworm. Environ. Entomol. 8:993-998.

Kitching, R. C. 1971. A simple simulation of dispersal of animals among units of discrete habitats. Oecologia. 7:95-116.

Leuis, K. 1%7. Influence of wild flowers on parasitism of tent caterpillar and codling moth. Canadian Entomol. 99:444446.

Lewis, F. B. 1960. Factors affecting assessment of parasitization by Apantelesfumiferanae and Glypra fumiferanae on spruce budworm larvae. Canadian Entomol. 92:881491.

McLintock, T. F. 1955. How damage to balsam fir develops after a spruce budwom epidemic. USDA Forest Sew. Sta. Pap. 75, Northeast Forest Exp. Sta., Upper Darby, Pa.

Miller, C. A. 1958. The measurement of spruce budwom populations and mortality during the first and second larval instars. Canadian J. Zool. 36:409-442.

Monis, R. F., and D. G. Mott. 1%3. Dispersal and the spruce budworm. p. 18CL189 in: R. F. Monis (ed.). The Dynamics of Epidemic Spruce Budworm Populations. Entomol. Soc. Canada Mem. 31: 1-332.


Mott, D. G. 1963. The forest and the spruce budworm. p. 189-201 in: R. F. Monis (ed.). The Dynamics of Epidemic Spruce Budworm Populations. Entomol. Soc. Canada Mem. 3 1:l- 332.

Price, P. W. 1976. Colonizations of crops by arthropods: non equilibrium communities in soybean fields. Environ. Entomol. 5:605410.

Shaw, J. G., and C. H. A. Little. 1973. Dispersal of second instar spruce budworm. Dept. Environ. Canadian Forest. Sew. Bimonthly Res. Notes. 29:3&31.

Simmons, G. A, , D. E. Leonard, and C. W. Chen. 1975. Influence of tree species and composition on parasitism of the spruce budworm, Chorisroneura furniferana (Clem.). Environ. Entomol. 4:832436.

Simmons, G. A., H. L . Brown and K. E. Gibbs. 1977. Environmental impact and efficacy of Imidan used for spruce budworm control in Maine. Maine Life Sci. and Agric. Exp. Sta. Misc. Rep. 189.

Syme, P. D. 1966. The effect of wild carrot on a common parasite of the European pine shoot moth. Canadian Dept. Forest., Bimonthly Res. Notes. 20:3.

. 1975. The effects of flowers on the longevity and fecundity of two native parasites of the European pine shoot moth in Ontario. Environ. Entomol. 4:336-346.

Turner, K. B. 1952. The relation of mortality of balsam fir caused by spruce budworm to forest composition in the Algoma Forest of Ontario. Canadian Dept. Agric. Pub. 875.

Watt, K. E. F. 1968. Ecology and Resource Management. McGraw-Hill, New York. Wellington, W. G. 1948. The light reactions of the spruce budworm, Choristoneura furni-

ferana (Clem). Canadian Entomol. 30:56-82. Williams, C. B. 1%6. Differential effects of the 1944-56 spruce budworm outbreak in eastern

Oregon. USDA Forest Sew. Res. Pap. PNW-33, Pacific Northwest Forest and Range Exp. Sta., Portland, Oregon.


FLEAS OF THE NORWAY RAT IN THE SUPERIOR, WISCONSIN, HARBOR AREA

(SIPHONAPTERA: CERATOPHYLLIDAE, HYSTRICHOPSYLLIDAE)

Darol L. ~ a u f m a n n ] and Norman D. ~ a d t k e ~

ABSTRACT

The occurrence of the fleas of the Norway rat, Rattus noruegicus (Berkenhout) was monitored in Superior, Wisconsin, for 12 months. A total of 441 fleas was recovered from 300 rat hosts. Nosopsyllus fasciatus (Bosc), the northern rat flea, was the most abundant species found (94.1%) while Ctenophthalmus pseudagyrtes pseudagyrtes Baker (5.4%) and Megabothris asio asio (Baker) (0.5%) were also present. The flea index was highest (4.8) in July and lowest (0.2) in February and March. Forty-four percent of the rats examined were infested.

Supenor, Wisconsin, is located at the western terminus of the Great Lakes and serves the upper Midwest as a grain storage and shipping center. The location of grain handling facilities in the port ensures the presence of a Norway rat population.

METHODS AND MATERIALS

Rats were trapped at two waterfront areas from June 1976 through May 1977. Trapping sites were located near railways in close proximity to grain elevators. Rat burrows and runways were found in debris-filled ditches overgrown with grass and brush. Grain spillage occurred on the rail facilities just short distances from these rat habitats.

Rats were trapped in Victor snap traps baited with a paste of sunflower seed and wheat flour. Trapped rats were removed within 10 minutes of capture and placed in heavy plastic bags for transport to the laboratory. Fleas were removed by employing a combination of blowing air against the grain of the fur, as described by Balthazard and Eftekhari (1957), and vigorous b r u s h over a white enamel pan. Fleas were preserved in 70% alcohol and later cleared in a lOnc solution of KOH.

RESULTS AND DISCUSSION

A total of 441 fleas was recovered from 300 rat hosts (25/month) from June 1976 through May 1977 (Table 1). The northern rat flea, Nosopsyllus fasciatus (Bosc) (Ceratophyllidae), was the most abundant species present. constituting 94.1% (415) of all fleas collected. Ctenophthalmus p s e u d a ~ r t e s pseudag~rtes Baker (Hystrichopsyllidae) and Megabofhris asio asio (Baker) (Ceratophyllidae) were also found, but in significantly lower numbers, making up 5 .45 (24) and 0.5Cr (2) of the fleas collected. N. fasciatus has been found on Rattus norregicus in Wisconsin on several occasions (Knipping et al. 1950. Haas and Dicke 1959, Haas and Wilson 1973). C . p. pseudagyrres and Megabofhris spp. occur on a wide variety of hosts. mostl\- microtine and cricetid rodents (Benton and Kelly 1969, Haas and Wilson 1973). C . p. pseudag?.rtes comprised 15V of the fleas collected from R . noruegicus in Kingston, R h d e Island (Knutson and Sqrnkowicz 1952). The occurrence of C . p. pseudagyrtes and M . a. asio on the S o m a y rat in Superior can possibly be explained by the presence of their more normal hosts in the collection areas.

l~e~anrnen t of Biolop. LniLersih of Wisconsin-Superior. Superior. WI 54880. 2~chool of Public HsalIh. L-niversity of California-Los Angeles, Los Angeles, CA 90024.


Table 1. Fleas recovered from 25 Norway rats examined per month from June 1976 through May 1977.

Number Nosopsyllus Ctenophthalrnus Megabothris Month of Fleas fusciatus p. pseudagyrtes asio asio

Jun 78 69 8 1 J ul 120 116 3 1 Aug 48 47 1 0 S ~ P 47 44 3 0 Oct 35 33 2 0 Nov 23 20 3 0 Dec 20 20 0 0 Jan 13 13 0 0 Feb 4 4 0 0 Mar 5 5 0 0 A P ~ 22 19 3 0 May 26 25 1 o

Totals 44 1 4 15 24 2

Table 2. Rat flea infestation rates for 25 Norway rats examined per month from June 1976 through May 1977.

No. (%)Rats Max. No. FleaslInfested Flea Month Infested FleasIRat Rat Index

Jun 21 (84) 11 3.7 3.1 Jul 24 (96) 13 5.0 4.8 Aug 13 (52) 17 3.7 1.9 S ~ P 14 (56) 11 3.4 1.9 Oct 9 (36) 8 3.9 1.4 Nov 9 (36) 9 2.6 0.9 Dec 5 (20) 8 4.0 0.8 Jan 8 (32) 5 1.6 0.5 Feb 3 (12) 2 1.3 0.2 Mar 4 (16) 2 1.3 0.2 A P ~ 10 (40) 6 2.2 0.8 May 13 (52) 4 2.0 1.0

Total (Mean) 133 (44) (3.3) (1.5)

The small sample of hosts in this study does not allow for a definitive judgement of seasonal distribution. The number of N. fasciatus colIected was greatest (116) in July and least (4) in February. N . fasciatus was taken in maximum numbers in winter and spring and in minimum numbers during Late summer in Baltimore, Maryland (Yeh and Davis 1950). At Norfolk, Virginia, N. fasciatus was found to be most prevalent in March (Hasseltine 1929). This species attained a peak population in the spring and again in the fall in Philadelphia, Pennsylvania (Vogel and Cadwallader 1935). An early spring population peak (March- April) and a late summer peak (July-August) for C. p. pseudagyrtes has been reported in northeastern U.S. (Benton and Kelly 1%9); very few rat hosts were involved, however.


Fleas were found on 133 (44%) of the rats examined with a maximum of 24 (96%) individuals infested in July and only 3 (12%) in February. The number of fleas on any single host was usually small although 17 were recovered from one rat in August. The average number of fleas per infested rat ranged from 1.3 in February and March to 5.0 in July with an average of 3.3 fleas found on each of the 133 infested hosts taken during the study. The flea index ranged from 0.2 in February and March to 4.8 in July with an average of 1.5 for the year (Table 2). Yeh and Davis (1950) found 54.5% of 966 Norway rats to be infested with fleas with an average of 6.7 fleas per infested rat. Other authors have noted infestations of from 35% to 41% with an average of 4.0 to 4.7 fleas per infested rat (Degiusti and Hartley 1965, Knutson and Szymkowicz 1952, Vogel and Cadwallader 1935). Flea indices determined by these same authors are similar to that found in this study.

ACKNOWLEDGMENT

We thank Dr. Nixon Wilson, Department of Biology, University of Northern Iowa, for his helpful suggestions and for his confirmation of the identification of the flea species involved in this study.

LITERATURE CITED

Balthazard, M. and M. Eftekhari. 1957. Techniques de recoite, de manipulation et d'elevage des puces de rongeurs. Bull. WHO. 16:436-440.

Benton, A. H. and D. L. Kelly. 1969. Notes on the biology of Ctenopl7rl7almus p. pseudagyrtes Baker in the Northeast (Siphonaptera: Hystrichopsyllidae). J. New York En- tomol. Soc. 77:70-74.

Degiusti, D. L. and C. F. Hartley. 1965. Ectoparasites of rats fromDetroit, Michigan. Amer. J. Trop. Med. and Hyg. 14:309-313.

Haas, G. E. and R. J. Dicke. 1959. Fleas collected from cottontail rabbits in Wisconsin. Trans. Wisconsin Acad. Sci., Arts, Letters. 48:125-133.

Haas, G. E. and N. Wilson. 1973. Siphonaptera of Wisconsin. Proc. Entomol. Soc. Wash- ington 75:302-3 14.

Hasseltine, H. H. 1929. Rat-flea survey of the port of Norfolk, Virginia. Pub. Health Rep. 44579-589.

Knipping, P. A., B. B. Morgan, and R. J. Dicke. 1950. Preliminary list of some fleas from Wisconsin. Trans. Wisconsin Acad. Sci., Arts, Letters. 40:199-206.

Knutson, H. and R. T. Szymkowicz. 1952. Ectoparasitism of Norway rats in an inland New England village and in a New England seaport. J . Econ. Entomol. 45:331339.

Vogel, C. W. and C. Cadwallader. 1935. Rat flea survey of the port of Philadelphia, Penn- sylvania. Pub. Health Rep. 50:952-957.

Yeh, J. and D. E. Davis. 1950. Seasonal changes in abundance of fleas on rats at Baltimore, Md. Pub. Health Rep. 65:337-342.

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