WWWWW WWWWW WW L— 205 THESIS 3 129300 .._—-___. A q - -. o‘q-uur' ' ' 7 r i . 4 "v .5 ‘J EP'.” 9-0 . _‘. ('I‘ no t .. Ar..,_.__ a, “-4 d5...-‘,,;..; I 2 .7 O. Q4 1...... oooooo i I This is to certify that the thesis entitled Activity patterns in some species of deciduous forest Litter Carabidae (CoLeoptera) in Michigan' 5 Upper Peninsula presente Judy Mousigian Nesmith has been accepted towards fulfillment ' . . of the requirements for M.S. degree in Zoology / /z/,‘ / professor Ri. a . . J. Snider mgr 07/ //f:> 0.7539 MS U is an Affirmative Action/Equal Opportunity Institution MSU RETURNING MATERIALS: Place in book drop to LJBRAfiJES remove this checkout from .—c—. your record. FINES will be charged if book is returned after the date stamped below. SEP ? . . WEB 6 1995 U, eoifil’S 1.x, ~11?” 1. l 1. ’ W r . V p . .t‘; 1‘ 54': L53", ‘2. ’ha amiss ACTIVITY PATTERNS IN SOME SPECIES OF DECIDUOUS FOREST LITTER CARABIDAE (COLEOPTERA) IN MICHIGAN'S UPPER PENINSULA by Judy Mousigisn Nesmith A Thesis Submitted to Michigan State University in partial fulfillment of the requirements for the degree of ‘ MASTER OF SCIENCE Department of Zoology 1985 05 ABSTRACT 3 ACTIVITY PATTERNS IN SOME SPECIES OF DECIDUOUS FOREST LITTER CARABIDAE (COLEOPTERA) IN MICHIGAN'S UPPER PENINSULA by Judy Mous igian Ne smith Pitfall traps were used to survey Carabidae (Coleoptera) in a deciduous woodlot in Michigan's Upper Peninsula. Five species of carabids were selected for analysis. Seasonal and dial activity patterns of Calathus ingratus Dejean, Pterostichus coracinus Newman, 2, melanarius Illiger, P: pensylvanicus Leconte, and Sunuchus impunctatus Say were assessed. Effects of temperature, trapping date and gender upon activity patterns were investigated for each species. The species studied displayed predominantly nocturnal behavior, though each had some incidence of diurnal activity. Temperature/activity relationships varied between species. Increases in overall activity corresponded closely with these species breeding seasons. In most species, diurnal activity and female activity peaked during periods of reproduction. The necessity for individuals, especially females, of each species to seek extra food, mates, and breeding and oviposition sites is thought to have contributed to increased carabid activity during breeding seasons in this study, and to have induced diurnalism in these normally nocturnal species. ii To my parents, in appreciation for their love and wisdom. iii ACKNOWLEDGEMENTS I wish to thank Dr. R. J. Snider for serving as chairman of my guidance committee. Thanks are also due to Drs. R. Fischer and M. M. Hensley who served as members of my guidance committee. I also wish to thank Gary Dunn of the MSU Cooperative Extension Service for verifying and correcting my identifications of carabid specimens. Special appreciation is extended to the Zoology Department office staff for their wonderful help and good humor through the years. Most of all, I wish to extend my deepest appreciation to my family and friends, and especially my husband Neron, for their love, support, and understanding. Table Table Table Table Table Table Table l. 2. 3. 4. 5. 7. 8. iv LIST OF TABLES List of carabid species collected by pitfall trapping in the study area with number of individuals of each species. . . . . . . . . . . . . Regression statistics of weekly night and day trap catches of the 5 most common species of Carabidae collected within the study area regressed on minimum and maximum temperatures in °C. . . . . . . Average number of carabids collected per hour at night and during the day for all species and for the 5 most common species at the study site. . . . . Results of analysis of variance of number of individuals collected per hour by diel factors for the 5 most common species at the research site . Product-moment correlation statistics of night: day trap collections of the 5 most common carabids investigated and their respective body lengths . . . Means and standard deviations of males and females caught per trap and results of comparisons using t or Q‘ tests. 0 s e e e s s e o s s o e s s 0‘. e e 0 Means and standard deviations of male:female ratios and results of comparisons between night and day catChas “Sing t Of q- tests. 0 e e e s s s s e s o o 16 25 39 39 43 45 47 Figure Figure Figure Figure _ Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure 1. 2. 3. 4. 5. 6. 7. 8. 9. ll. 12. 15. V LIST OF FIGURES Dialnson County, mChigan e e e a e e e s e e Location of study area . . . . . . . . . . . . Diagram of pitfall trap. . . . . . . . . . . . Minimum and maximum daily temperatures recorded at the way Dam weather station from May, 1983 through OCtOber , 1983 O O O O O O O O O O O O 0 Number of male, female and teneral individuals of E. melanarius caught per trapping date. . . Number of male, female and teneral individuals of g, coracinus caught per trapping date . . . Number of male, female and teneral individuals of S. impunctatus caught per trapping date . . Number of male, female and teneral individuals of g. ingratus caught per trapping date. . . . Number of male, female and teneral individuals of g, pensylvanicus caught per trapping date . Number of individuals of g, coracinus caught in diel traps in spring, summer and fall, plotted against temperature in .C. . . . . . . Number of individuals of _S_. impunctatus caught in diel traps in spring, summer and fall, plotted against temperature in °C. . . . . . . Number of individuals of g, melanarius caught in diel traps in spring, summer and fall, plotted against temperature in 'C. . . . . . . Number of individuals of g. ingratus caught in diel traps in spring, summer and fall, plotted against temperature in “C. . . . . . . Number of individuals of g, pensylvanicus caught in diel traps in spring, summer and fall, plotted against temperature in °C. . . . . . . Number of male, female and teneral individuals of g, impunctatus caught per hour at night and during the day for each trapping date. . . . . 10 12 18 19 20 21 22 26 27 28 29 30 34 Figure 16. Figure 17. Figure 18. Figure 19. vi LIST OF FIGURES (Continued) Number of male, female and teneral individuals of £3 melanarius caught per hour at night and during the day for each trapping date. . . . . . Nmmber of male, female and teneral individuals of g, coracinus caught per hour at night and during the day for each trapping date. . . . . . Number of male, female and teneral individuals of E, iggratus caught per hour at night and during the day for each trapping date. . . . . . Number of male, female and teneral individuals of g, pensylvanicus caught per hour at night and during the day for each trapping date. . . . . . 35 36 37 38 vii TABLE OF CONTENTS DEDICATION . . . . . . . . . . . . . . . . . . . . . . . . . . iv ACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . .iii LIST OF TABLES . . . . . . . . . . . . . . . . . . . . . . . . iv LIST OF FIGURES. . . . . . . . . . . . . . . . . . . . . . . . v INTRODUCTION AND LITERATURE REVIEW . . . . . . . . . . . . . . 1 MATERIALS AND METHODS Experimental Area . . . . . . . . . . . . . . . . . . . . 5 Sanpling.........................8 Sorting . . . . . . . . . . . . . . . . . . . . . . . . . 11 Heather Data. . . . . . . . . . . . . . . . . . . . . . . 11 Statistical Analysis. . . . . . . . . . . . . . . . . . . 13 RESULTS AND DISCUSSION General Results . . . . . . . . . . . . . . . . . . . . . 15 Seasonal Activity . . . . . . . . . . . . . . . . . . . . 15 Temperatureufec:ssoeeoss00000000000024 Diel Activity . . . . . . . . . . . . .g. . . . . . . . . 33 “we: Effect.. 0 O O O 0 O O 0 O O O O O O O O O O O O O 44 smryOOOOOOOO0.00.0.000000000048 APPENDICES A. Formulae for Vegetation Analysis. . . . . . . . . . . 50 E. Trapping Schedule . . . . . . . . . . . . . . . . . . 51 C. Minimum and maximum.temperatures used for regression statistics in °C . . . . . . . . . . . . . 52 D. Average night and day relative humidities. . . . . . 53 E. Length, in hours, of night and day trapping periods through the season. . . . . . . . . . . . . . 54 F. Night/day sample data . . . . . . . . . . . . . . . . 55 G. Twenty-four hour sample data. . . . . . . . . . . . . 59 LITERATURE CITED INTRODUCTION AND LITERATURE REVIEW Rhythmic periodicity is prevalent throughout the animal kingdom (Calhoun, 1944). Proper timing of activity in insects is of particular importance. Appearance of adults and larvae must frequently coincide with availability of suitable food supplies and reproduction must be synchronized correctly to ensure that resistant life stages are present during unfavorable seasonal conditions. Carabid beetles, like many insects, show both daily and seasonal patterns of activity. Among the first to document periodicity in carabid beetles were Park and Keller (1932) and Krumbiegel (1932). Since then, under both laboratory and field conditions, rhythmic activity patterns have been reported in carabids by numerous investigators (Calhoun, 1944; Williams, 1959; Greenslade, 1963; Grum, 1966; Breymeyer, l966a,b; Thiele and Heber, 1968; Kirk, 1971b; Dondale et al., 1972; Luff, 1978; Bears, 1979, Doiteau, 1983; Boucher and Malausa, 1984). Various techniques have been used to assess carabid activity patterns. Laboratory research has been conducted using both direct observations and actographs under natural and artificial light-dark regimes (Williams, 1959; Grum, 1966; Thiele, l977a,b; Boucher and Malausa, 1984). Actographs and observations of beetles in enclosures have also been used in field studies (Greenslade, 1963; Dennison and Hodkinson, l984b). Kirk (1971b) observed beetles in their natural habitat. Finally, Baars (1979) captured, marked, and radioactively tagged beetles, then followed their movements after release at their capture site. The majority of carabid activity studies were assessed by pitfall trapping. A few investigators have used time-sorting pitfall traps (Williams, 1959; Dondale et al., 1972; Luff, 1978; Stubbe et al., 1984). These traps have a clock-driven mechanism to partition catches by time increments. Most researchers, however, use a buried trap design (Greenslade, 1965; Barlow, 1970; Jones, 1979; Levesque et al., 1979; Desender and Maelfait, 1982; Dennison and Eodkinson, 1984a,b). Definite diel patterns of activity were reported for many of the species studied. Greenslade (1963) classified diel activity into three well-defined patterns - nocturnal, diurnal, and plastic. Later, Thiele and Heber (1968) summarized diel activity among carabid beetles in a comprehensive literature review. They concurred with Greenslade's proposed classification, and reported that, of the species investigated, 60! were strictly nocturnal, 222 were mainly day-active, and 182 showed intermediate behavior. Plastic, or intermediate carabid beetles exhibited varying degrees of diurnalism or nocturnalism in response to different environmental conditions. Frequently, seasonal reproductive rhythms are associated with diel activity patterns (Thiele and Weber, 1968). Thiele (1977c) reports that Larsson (1939) and Lindroth (1949) were among the first researchers to recognize different types of carabid seasonality. Greenslade (1965) simplified their definitions and divided carabid seasonality into three types: 1) spring breeders, 2) summer and summer-autumn breeders, and 3) autumn breeders. Subsequent research has revealed annual reproductive and activity. patterns for many species (Johnson et al., 1966; Barlow, 1970; Kirk, l97la,b; Levesque et al., 1979; Desender et al., 1981; Desender, 1983; and Dennison and Hodkinson, 1984a). Paarman (1979) later expanded Greenslade's categories to include the annual rhythms of species from tropic and sub-tropic areas. Despite previous studies, mechanisms regulating rhythmic activity patterns are not clearly understood. Activity patterns are most often manifestations of an endogenous periodicity (Saunders, 1982). Frequently, other factors override autonomous control and readjust patterns of activity (Beck, 1980; Saunders, 1982). Rythmicity in carabid beetles is particularly malleable by other factors, and varies widely between species (Thiele and Haber, 1968; Thiele, 1977c). Carabid beetles in Michigan's mostly forested Upper Peninsula are poorly known, especially in its western half. The only surveys conducted in this area were by Hubbard and Schwarz (1878) and Andrews (1921). Neither were extensive and few carabids were reported. It is the purpose of the following study to supplement the distributional records of Carabidae in Michigan, and contribute to the limited available knowledge of the habits of North American forest carabids. In summary, the following study: 1. surveyed using pitfall traps, and identified the surface-active Carabidae in a deciduous woodlot in Dickinson County, Michigan. 2. documented the diel activity patterns of the most common carabid species encountered. 3. investigated the relationship between activity patterns, temperature, and trapping date. 4. examined differences in activity patterns between species and between sexes of the same species. This study was part of an ecological monitoring program for the United States Department of Defense' Extremely Low Frequency (ELF) Communications System. Portions of this program.were subcontracted to Michigan State University from IIT Research Institute, which, in turn, was contracted by United States Naval Defense. This study was included in task 5.3 - Soil and Litter Arthropoda. MATERIALS AND METHODS EXPERIMENTAL AREA This study was conducted on state land in the western end of Michigan's Upper Peninsula, T43N, R29W, 88 (Figures 1,2). The site encompassed about two acres and was approximately seven miles east of the town of Channing in Dickinson County. This area is part of the Canadian shield, a region of the Upper Peninsula where strong precambrian bedrock is close to the surface and the topography is typically characterized by high hills and deep valleys. Glaciation has left a layer of glacial till on top of this bedrock (Dorr & Eschman, 1970). The soil at the study site was thin, undeveloped below 20 inches, and rocky, moderately well to well-drained, with a humus layer of only 2-3 inches. It typed out as an Emmet fine sandy loaml. The area was once covered by dense white and red pine forest. However, subsequent logging, which reached its peak in the 1890s, has eliminated this forest association (Santer, 1977). The study site was last logged between 1964 and 1969 and is now a mature hardwood forestz. Tree density at the site was approximately 11 trees per 100m2. Sugar maple (A555,saccharum.Marsha11) was by far the most prevalent tree species, but the largest trees at the site were 1,2. The author is indebted to Halt Summers of the 0.8. Soil Conservation Service in Iron Mountain, MI, for providing information about the soil and history of the study area. Figure 1. Dickinson County, Michigan. 6 marks research site. SCALE IN Mme ruu- 220/ ' ‘ Figure 2. Location of study area. basswood Qgilia_americana Linnaeus). Cherry (ngggg_serotina Ehrhart), American hop-hornbeam (925523 Virginians (Miller) K. Koch), paper birch (gagglg’papyrifera (March)) and American elm (91223 americana (Linnaeus)) also occurred at the site but were few in number. The understory layer at the study site was sparse, with approximately 62 coverage, and consisted of two shrub species, leatherwood (Dirge palustris Linnaeus) and hazelnut (Corzlus sp.). Herb cover was moderate, covering abut 281 of the study area, and, out of 20 different plant types, sugar maple seedlings, sedges (Cyperaceae), wild lily of the valley (Maianthemum canadense Desfontaines), and sweet cicely (Osmorhiza claztoni (Clarke) were most prevalent. Yellow violet (giQISDEubescens Aiton), American hop-hornbeam seedlings, and spikeweed (Aralia racemosa Linnaeus) were also common. The formulae used to calculate plant density or coverage are given in Appendix A. SAMPLING Samples were taken using 40 randomly placed pitfall traps, 20 in each of two adjacent areas at the study site. Sampling periods encompassed twenty-four hours. Half of the traps were used to collect separate night and day samples. These traps were set at dusk and changed and refilled the following dawn. The second sample was collected at dusk the same day. The remaining traps were set at dusk and collected at dusk the next day. Traps were covered between sampling periods. Samples were collected in this manner weekly from May through October, 1983, for a total of twenty-five sampling dates. No sample was taken the third week of October due to severe weather. The sampling schedule is provided in appendix 3. Adis (1979) summarized the problems associated with pitfall trapping for arthropods and suggested that pitfall trap results should be interpreted with caution. Problems arise in separating the density and activity components of a given population. However, Briggs (1961) and Greenslade (1964) found that pitfall trapping was useful in studies of carabid beetles for assessing strictly locomotor activity. Thiele (1977c) reports that for analysis of daily and annual activity rhythms, pitfall trapping is superior to any other method. Further studies have shown that pitfall trapping may be the best method for investigating carabid community composition. Kirk (1971b) compared pitfall trapping to soil sampling and direct observation in cropland species. Pitfall trapping produced the same number of species in similar ratios of abundance as the other sampling methods. Uetz and Unzicker (1976) investigated pitfall trapping and quadrat litter sampling efficiencies in assessing populations of woodland cursorial spiders. Dennison and Hodkinson (1984b) conducted similar research on carabid and staphylinid beetles. Both studies showed that pitfall trapping provided a more complete faunal list of cursorial arthrOpods than quadrat sampling. The pitfall traps used in this study were made of smooth white plastic and consisted of three parts; a large cup which functioned as an outer sheath, a smaller cup which fitted snugly inside the large cup, and a funnel that rested on the rim of the large cup (figure 3). The funnel was used to help retain trapped beetles in the small inner cup, and to reduce incidence of disruption by other animals. Holes were punched into the bottom of the large cup and 8.5 cm.---'9 AN. Figure 3. Diagram of Pitfall Trap sides of the small cup for drainage. Clear plastic cups upended over the pitfall traps served as covers when the traps were not in use. Traps were buried with rims flush with the soil surface. They were then covered and left undisturbed for a week. This served to eliminate ”digging-in effect“, a phenomenon in which disturbance and raised 002 levels in the soil caused by placing traps create artificially high levels of arthropod activity for the following two days (Joosse and Kapteijn, 1968). Traps were set by filling the inner cup with approximately 2.5 centimeters of ethylene glycol, which acted as a trapping agent and temporary preservative. Changing traps was simply a matter of replacing the inner cup with a fresh cup and refilling with ethylene glycol. ll SORTING A solution of 951 ethyl alcohol with 12 glycerin was used to rinse samples through a fine mesh sieve (U.S. Standard sieve series #2000) and for subsequent storage. Samples were later sorted and carabids were pinned and identified to species using Lindroth's (1969) system of classification. Sex and stage of adult (teneral or mature) were also identified and recorded along with number of individuals. Teneral adults were recognized by their pale, soft exoskeleton. Adult carabids generally remain teneral for approximately 2-3 days after pupal eclosion (Goulet, 1974). Voucher specimens are deposited in the Entomology Museum of Michigan State University. WEATHER DATA Temperature data were obtained from the National Weather service. Minimum and maximum daily temperatures were recorded at the Way Dam weather station situated approximately twelve miles west of the study area (figure 4). Minimum daily temperatures for night catch regressions were obtained by averaging minimum temperatures of the two adjacent dates of the trapping period. For example, minimum.daily temperatures of May 4th and 5th were averaged to obtain the minimum temperature used for the night catch of that sampling date. Minimum and maximum temperatures used for regression analyses are listed in appendix C. Hmmidity data were not available from the Way Dam.weather station. A few weeks of humidity data, however, were obtained from a hygrothermograph situated at the site which functioned intermittently. These data are tabled in appendix D. 12 .mwm~ .umoouoo :wsousu mama .mmz Boom moaumum nonumoa Eon has ecu um ooeuoumu mousumumnaou Adamo asawxma use snags“: .q ouswfim Foo mum o3< . one 235 on m. on m. on m. on . m... on m. on b b h P I b l3 STATISTICAL ANALYSIS Analyses of variance (using an ANOVA computer program SPSS version 8.3, 1984), linear regression analyses, and T- and q- tests of comparisons were used to compare catch size with gender, date, diel effects, and temperature. Only species caught in sufficient numbers for these analyses (52 of more of the total catch) were examined. For statistics comparing diel effects, numbers caught per sample were converted to numbers caught per hour to account for the changing lengths of day and night through the season. A list of length of night and day trapping periods is presented in appendix E. Data for the ANOVA statistics were log-transformed so that they would be approximately normally distributed. The remaining statistics were hand-calculated using procedures and formulae from Sokal and Rohlf (1981). Accepted significance for statistical results was p - 0.05 or less. Numbers per sample and numbers caught per hour were separated into male/female and night/day components for each species (appendices F and G). Seasonal means were calculated and compared using T- and q-tests. Number of individuals and numbers caught per hour were plotted against trapping date to display shifts in diel and seasonal activity. Occurrence of teneral adults was indicated to identify periods of recruitment. Catches per trapping date were regressed on temperature for that date for each species. Night catches were regressed on minimum night temperatures and day catches regressed on maximum day temperatures based on Bear's (1979) conclusions that minimum temperatures were significant to night-active species, while day 14 active species were influenced more by maximum temperatures. Catches for each species were analysed in two-month sections (n=9 in the spring months, n=8 in each of the remaining sections). This procedure served to reduce the confounding effects of seasonal influences on activity. 15 RESULTS AND DISCUSSION GENERAL RESULTS Eighteen different species of Carabidae were collected from May 1983 through October 1983. Examination of museum records showed that ten species had not previously been recorded as occurring in Dickinson County. A list of these species is provided in Table 1. Inspection of catch data showed five species caught in far greater numbers than the others. Calathus ingratus Dejean, Pterostichus coracinus Newman, 2, melanarius Illiger, g. penaylvanicus Leconte, and Szguchus impunctatus Say each contributed over 52 to the total catch. Individuals of 2:. pensylvanicus alone made up over a third of the total catch. Together, these species comprised over 901 of all carabids trapped. These species were selected for further analysis. All selected species belong to the tribe Pterostichini and are commonly found in North American forest habitats. ‘2, melanarius is an introduced European species and is also common in fields and arable land (Lindroth, 1969; Barlow, 1970; Ericson, 1977; Jones, 1979). The remaining species are native to North America (Lindroth, 1969). Activity in each of these selected species as determined by number trapped, was assessed and compared with a variety of factors. Data on these carabid beetles activity patterns, some possible influences on activity, and variation of activity between 'species will be discussed in the following pages. SEASONAL ACTIVTTV Each carabid species examined displayed a distinct seasonal 16 Table 1. List of carabid species collected by pitfall trapping in the study area with number of individuals of each species. An * denotes species that are new records for Dickinson County. species number of individuals caught ‘Agggg! decentis Say 1 *A. placidum Say 1 A. retractum Leconte l9 *Bembidion quadrimaculatum oppositum Say 1 Calathus gregarius Say 57 *CE ingratus Dejean 206 Clivina £2222£_Linne 3 *ngindis cribricollis Dejean 9 Hagpalus fuliginosus Duftschmid 24 Mugg'cyanescens Dejean 12 *Notiophilus £32222_Herbst 3 *Pterostichus adstrictus Eschscholtz 6 z- m 38? 3 fig, coracinus Newman 131 g, melanarius Illiger 108 E? pensylvanicus Leconte 526 *Sphaeroderus lecontei Dejean 1 *Szggchus impunctatus Say 272 TOTAL 1383 i of species 18 l7 pattern of activity during 1983 (Figures 5-9). Occurrence of teneral adults indicated periods of recruitment (e.g. additions of new adults to the adult population) for each species. Seasonal data for g, melanarius and g, coracinus showed activity beginning in May, gradually increasing during the summer months, and declining in September. Activity of P. melanarius lagged a few weeks behind that of £3 coracinus (Figures 5,6). Teneral adults of both species were caught in July, several weeks after activity began. S, impunctatus also was most active during summer. Activity in this species, however, started in late June, rose sharply to a peak in July, and ended by early September (Figure 7). Tenerals made up the entire first catch in June. Activity of another summer-active species, 9, iggratus, began in mid-June, peaked during August, and continued through late October (Figure 8). Tenerals first appeared in late July, but most were caught in August. Seasonal activity of £3 pensylvanicus differed greatly from the other species studied (Figure 9). The majority of g. penaylvanicus were caught in May, June and July, and declined abruptly in August. A secondary activity peak occurred in autumn. Teneral adults were present only during this second peak. The seasonal activity patterns shown by the species in this , study corresponded closely with patterns and information documented in previous investigations (Briggs, 1961); Greenslade, 1965; Barlow 1970; Levesque et a1, 1979; Jones, 1979). Although onset of activity may have varied by a few weeks depending upon temperature 18 msfiumamaoe . a .2221m Ko 3.2.3-8 .oomo wofimuouo Hog unmade Im.mo mamsefi>woma Houooou one mamaom .oama wo Homauz .m unawam 03 .51. 22. >32 «.0 c.-- 0. 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L o._ . g L Q. . .22... «-m “-0 0.? >t>r6< 50.2 .9362 Mul§ E... 39 Table 3. Average number of carabids collected per hour at night and during the day for all species and for the 5 most common species at the study site. species 2. ingratus £9 coracinus g. nelsnar ius 'U o pensylvanicus _S_. igpunctatus All Carabids night catch/hour day catch/hour .49 .02 .29 .04 .25 .08 .72 .19 .36 .06 2.34 9.49 Table 4. Results of analysis of variance of number of individuals collected per hour by diel factors for the 5 most common species at the research site. hour were log-transformed. significance at P5 .001. Species '2“ pensylvanicus melanarius ["U o '2‘ coracinus '5“ inpunctatus '9 ingratus Source of variation diel diel diel diel diel Numbers collected per An * denotes accepted 3mm of Degrees of Mean squares freedom 3.093 1 .137 1 1.972 1 2.959 1 2.469 1 square 3.093 .137 1.972 2.959 2.469 F statistic 17.908* 1.863 38.150* 34.428* 18.303* 40 The probability that diel effects influenced activity in g, melanarius was 822. This species was also predominantly night-active, but the difference between night and day activity was not as great as other species exmained (Table 3). This study's results confirm conclusions reached by previous authors, nocturnal behavior was documented in g. melanarius in European studies (Greenslade, 1963; Luff, 1978; Dennison and Hodkinson, 1983), and was recorded in limited observations of 23' coracinus, £3 pensylvanicus, and g, igpunctatus in eastern Canada (Levesque et a1, 1979). Although carabid activity in this study primarily occurred at night, each species had some incidence of diurnal activity. Levels of this activity varied between species, but except in the case of 99 ingratus, a species that showed very little diurnal activity in this study, seasonal occurrences of diurnal activity in the species investigated generally followed their overall seasonal pattern of activity (Figures 15-19). Diurnal activity in these species tended to increase during late spring and summer when their overall activity increased. Diurnal activity in g. ingratus, on the other hand, also occurred near the beginning and end of this species' season of activity (Figure 18). 4 In field experiments in England, Luff (1978), and Dennison and Hodkinson (1983) found that small carabid species tended to show more diurnal activity than larger species. Their conclusions were based on negative correlations between day activity and mean body size. Luff (1978) suggested that larger species, being more conspicuous during the day, must conceal themselves from predators 41 by hiding deep in litter. Small species, on the other hand, are readily overlooked and can easily find shelter. In order to test Luff's theory, the product-moment correlation coefficient was calculated for the log of the ratio of mean night and day catches per hour and mean body size for each species. Results refuted Luff's hypothesis (table 5). The correlation coefficient was not significant, but its negative value indicated a trend towards negative correlation between body size and nocturnalism. Netably, Q, ingratus, the smallest species studied, was the least diurnally active of the five species investigated, and g, melanarius, a large species, was the most diurnally active. Luff's (1978) predation theory does not seem to apply in this study. Many authors theorize that nocturnal species of carabids, especially those in forest habitats, are particularly susceptible to desiccation (Kirk, 1971b; Greenslade, 1965; Thiele and Haber, 1968; and Thiele, 1977c). Restricting their activity to night reduces the exposure of these species to dry conditions since daytime relative humidities are often fairly low, rising only at night (Thiele and Haber, 1968; Bauer, 1977). Relative humidities in this study, in fact, regularly rose at night and declined substantially during the day (Appendix D). It is suggested that changing humidity levels were a contributing influence on nocturnal activity in the species examined. The tendency in this study towards diurnal activity in g. melanarius, a species found in field as well as forests (Lindroth, 1969), may have been due to a greater ability of this species to withstand drier air than the other carabids in this study. Most field 42 species of carabids are adapted to tolerate dry field conditions and hence are not as averse to diurnal activity as forest species (Thiele and Haber, 1968). Additionally, field notes recorded daytime rain during the few isolated instances that g, ingratus was caught in day traps over the season. This species appeared to be active almost solely during periods of high relative humidity and may be particularly sensitive to desiccation. Table 5. Product-moment correlation statistics of nightzday trap collections of the 5 most common carabids investigated and their respective body lengths. species log average #Ihour-nighg’ mean body length average #lhour-day (mm) 9. ingratus 1.39 7.7 i .90 _S_. impunctatus .78 9.5 i .67 g. pensylvanicus .58 11.5 1 .50 _P_. coracinus .86 16.6 i .80 g. melanarius .49 17.0 i .89 Some authors suspect that carabid activity patterns are primarily determined by competition. Williams (1959) and Breymeyer (1966) both postulated that activity of potentially competing ground macrofauna, specifically, carabid beetles, in ”older", more evolved communities (e.g., forests) is more symmetrically arranged 43 around a twenty-four hour cycle due to previous competitive interactions than activity in ”younger", less evolved communities (e.g., fields) where competition has not yet forced species into different temporal niches. Though the species examined in this study were in an ”older” community as defined by Williams (1959), a mature hardwood forest, activity of each species occurred primarily at night, displaying an asymmetrical arrangement of activity. Hence, competition between species did not appear to regulate periodicity in this study. ' Intraspecific competition, however, may have operated on some of the carabid species investigated here. Increased diurnal activity in species examined in this study coincided with these species breeding seasons, a time when breeding adults were seeking mates and breeding as well as food (Barlow, 1970). This phenomenon was especially apparent in g. pensylvanicus. A comparison of this species spring and fall activity peaks showed that this species was much more day active during spring, its main period of reproduction. Though increased activity may have been stimulated by reproductive factors, competition between individuals of the same species may have also influenced this shift to diurnal activity in the normally nocturnal species of this study. GENDER EFFECTS Seasonal and dial activity was found to vary between sexes as well as between species. Comparison of means of males and females caught per trap revealed significant differences in activity between sexes in g, pensylvanicus,‘§. impunctatus and g, ingratus (Table 7). Females appeared to be more active than males for each 44 Table 7. Means and standard deviations of males and females caught per trap and results of comparisons using t or q- tests. * denotes accepted significance (p 5_.05). The q statistic was used when variances were unequal but coefficients of variation were equal. Species Mean Standard T or q deviation statistic E. pensylvanicus Overall males 5.10 2.49 4.56* females 8.02 3.19 spring males 2.60 1.70 7.23* females 4.30 2.37 fall males 1.82 1.51 1.92* females 1.20 1.36 _P_. melanarius males 1.55 1.48 .85 females 1.22 1.94 _P_. coracinus males 1.58 1.34 .38 females 1.70 1.47 _S_. imminctatus males 2.05 1.96 10.88* females 4.75 2.89 g. iggratus males 1.55 1.56 10.56* females 3.60 2.45 45 of these species. Admittedly, increased female catches in this study could have been interpreted as greater female density as well as increased female activity. However, it is unlikely that density alone accounted for the disparity between male and female catches because pitfall traps encompass both activity and abundance components in a population. Sex ratios of night and day samples were also compared for each species investigated in this study. Results showed that proportions of females and males significantly differed between night and day for all but one species (table 8). These differences most likely reflected changes in activity between sexes since relative densities of males and females would probably not change appreciably in the course of one day. Changes in sex ratios indicated that higher levels of femalezmale activity occurred at night compared to day for g, coracinus,‘§. impunctatus, and Q. ingratus. Ratios of female to male activity increased during the day for g. pensylvanicus rather than at night. Moreover, comparisons of night and day sex ratios during the spring and fall activity peaks of g. pensylvanicus revealed a shift in female activity over the trapping season. Diurnal proportions of females were significantly greater than those of males during the spring for this species, but autumn sex ratios were nearly equal between night and day. This study has shown a positive association between warm day temperatures and diurnal activity of 2, pensylvanicus during its spring breeding season. warm day temperatures may have particularly influenced breeding females in this species. Though adults are cold-tolerant, warmth is important for stimulation of reproduction and egg 46 Table 8. Means and standard deviations of male:fema1e ratios and results of comparisons between night and day catches using t or q- tests. The q statistic was used when variances were unequal but coefficients of variation were equal. Species Mean Standard T or q deviation statistic _11. melanarius female:male- night 1.17 1.03 .78 female:male- day 1.42 .99 .11. coracinus female:male- night 1.75 1.48 4.29* female:male- day 1.07 .77 g. impunctatus female:male- night 2.34 2.05 5.22* female:male- day 1.72 .83 g. ingratus female:male- night 1.86 1.20 14.15* female:male- day 1.12 .39 _11. pensjlvanicus Overall female:male- night 1.25 .67 7.72* female:male- day 2.26 1.36 spring female:male- night 1.43 1.01 2.16* female:male- day 2.26 1.39 fall female:male- night .97 .56 .58 female:male- day 1.08 .60 47 development (Goulet, 1974; Thiele, 1977c). Increased female activity may be related to reproduction in the other species examined in this study. In addition to seeking mates many females must later select specific oviposition sites (Thiele, 1977c). Goulet (1974) found that females of g. pensylvanicus consistently chose the wettest areas of their habitat for oviposition. Although the oviposition habits of g. impunctatus, Q, ingratus, and Z: coracinus are poorly known, elaborate nesting behavior has been documented almost solely in the tribe Pterostichini, to which these species belong (Thiele, 1977c). Furthermore, gravid females may need extra nutrition for their developing eggs. Females trapped in this study tended to be larger than their male counterparts. SUMMARY Definite seasonal and diel activity patterns were revealed in g. pensjlvanicus, 1:. coracinus, g. melanarius, _S_. impunctatus and £9 ingratus in Michigan's Upper Peninsula. Seasonal patterns displayed by these species corresponded closely to their breeding seasons as documented in the literature. Increases in activity tended to coincide with periods of reproduction. Temperature was associated with activity in some species investigated. Night activity was positively associated with minimum temperatures in most cases, other species showed little or no response to temperature. '2, pensylvanicus, in particular, showed a shift in temperature relationships with activity through the trapping season. Each species was nocturnal. High humidity levels at night may have favored nocturnal activity in the species studied. Varying 48 amounts of diurnal activity, however, occurred in each species. In most species, this diurnal activity increased during breeding season. Intraspecific competition between breeding adults for mates and breeding areas may have induced diurnal activity in these normally nocturnal species. Females were more frequently captured than males in this study. Furthermore, activity levels of females varied disproportionately between night and day catches for some species. 2, pensylvanicus, again, showed seasonal shifts in female:male activity ratios. Increased female activity occurred during breeding seasons for most species investigated. Gravid females seeking food and oviposition sites may have contributed to this increase in activity. 49 >mwwzcnx > wonacpme non enmenmnpos asswwmom anew mans cosm»m%\wcc l~ I woo l~\am I e on «nose on penance > assocsnonom x essepne non sup «name e on names manocsnonee compasses on ocean I xnwmnpes mosuunw on evenne- > N >conmne comma mass on smashes > non newness > manna eons >veowcno noeonmmo on success > I nonsp wmmmnr on «nauseous panonomMnom rM enmepos > nonmp woman: on nauseonnm muc.acc any revenues no I swuowcne noesnmnw on enenpee > nonsp sauce oceansms >cuo~cno unencssnw on avenues > I e on nnmsmoORe absence > oceans an x wco nonsu u on nussseons AnOV revenues nnnacsanw on emeopse > I scoopcne mnensssmu on meanne- > x poo eel on sveowcne mnemsosonss non app evenne- colpzmsos on smashes > I «avenues assesses. on sweeps. >V+ surnames noemnsme on smashes > some mans >vso~sna unseceaow on newness > I e on muons encaps- > onosne n: x woe non-p u on muons A newanpes manganese on avenues > I scsowcno nuancesmn on mesons. > x —oo s:- nonsw on scsopcne mnsasosnpes >Veo~cno oocen on sweeps. > I >censmo noses «muses on escapes > ocon ~c uponm wepsnpee scene on avenues ? I > can upon I ocean sauce on meeapsm > an upon a a —co nonmp oocsn causes an upon n compasses on avenues > I announce mnoacesnw on scoops. > + meusnneo ocean on oneness > 50 >wwmzuHx a Hubwwmzn mommccrm «mu 2: 26 26 z: p n w a u a u m o po pp pn pw pm pm pa po p p no np nn no no no no nu no no oc up as: 26 zu as 26 2: p n u a m a u m o pa pp pn pw pm pm po pu pm po no np nn nu n» no no nu no no we pap . z: 2: 26 zu p n u e u o u a o po pp pn pu pa pw pm pu pa po no np nn no no no no nu no no no up ti 2: 26 ac 2U p n w a m w u m o pc pp pn pu pa pm po pu pm po nc np nn no n» no no nu no no we up m... U 2: 26 zu 2: p n u e u e u m o po pp pn pw pm pm pa pu pm po nc np nn nu n» nu no nu no no no can 25 26 . 25 p n u a u o u m o po pp pn pw pa pm p¢ pu pm po no np nn no no no no nu no no we up 51 APPENDIX C Minimum.and Maximum.temperatures used for regression statistics in ’0 date ‘ Maximum daily Averaged minimum temperature daily temperature 5/May 6.5 -4.0 12/May 24.0 1.5 19/May 20.5 3.0 26/May 9.0 0.5 2/Jun 21.0 2.0 10/Jun 21.5 4.0 17/Jun 18.5 7.0 23/Jun 30.5 17.0 30/Jun 24.0 8.5 7/Ju1 24.0 4.5 l4/Jul 31.5 13.0 Zl/Jul 33.5 17.5 28/Ju1 30.5 13.0 4/Aug 28.5 15.5 11/Aug 15.5 11.0 18/Aug 27.0 15.0 25/Aug 27.0 9.5 l/Sep 28.0 8.5 8/Sep 22.0 6.0 l4/Sep 16.5 0.0 22/Sep 10.0 2.0 29/Sep 24.5 12.0 6/0ct 17.0 6.5 13/0ct 5.0 1.5 27/0ct 11.5 2.0 52 APPENDIX D Average night and day relative humidities date average R.H.-night average R.H.-day 26/Aug 96.22 85.22 27/Aug 93.5 72.5 28/Aug 93.2 77.7 29/Aug 92.5 83.8 30/Aug 96.8 83.5 31/Aug 93.5 70.0 l/Sep 90.3 73.7 22/Sep 92.0 84.0 23/Sep 90.3 75.8 24/Scp 85.5 68.7 25/Sep 89.2 78.7 26/Sep 88.2 87.7 27/Sep 89.8 85.2 53 APPENDIX E Length, in hours, of night and day trapping periods through the eases date night length day length 4-5/May 9.25 15.25 11-12/Msy 8.50 15.00 18-19/May 8.50 15.00 25-26/May 8.00 16.00 1-2/Jun 8.00 15.50 9-10/Jun 8.00 16.00 16-17/Jun 8.25 17.00 22-23/Jun 8.00 16.00 29-30/Jun 8.25 15.50 6-7/Ju1 7.75 16.00 13-14/Jul 8.00 16.00 20-21/Ju1 8.25 16.00 27-28/Ju1 8.50 15.75 3-4/Ang 8.25 15.75 lO-ll/Aug 8.75 15.00 17-18/Ang 9.00 15.00 24-25/An8 9.25 14.85 319Ang/1-Sep 9.75 14.50 7-8/Sep 10.00 13.75 13-14/3ep 10.50 13.50 21-22/Sep 11.25 12.75 28-29/Sep 11.25 12.50 5-6/0ct 12.00 12.00 12-13/0ct 12.50 11.50 26-27/0ct 13.00 11.00 NIGHT/DAY SAMPLE DATA 9. ingratus 4-5/May ll-lZ/May 18-19/May 25-26/May 1-2/Jun 9-10/Jun 16-17/Jun 22-23/Jun 29-30/Jun 6-7/Ju1 13-14/Ju1 20-21/Ju1 27-28/Ju1 3-4/Aug lO-ll/Aug 17-18/Ans 24-25/Ang 31-Aug/1-Sep 7-8/Sep l3-l4/Sep 21-22/Sep 28-29/Sep 5-6/0ct 12-13/0ct 26-27/0ct TOTALS 0°0UH°0HHUINUJNHNON00000000 U U 00000000 .24 .25 .12 .82 .36 .80 .56 .11 .10 .04 .27 N !L_ hr 0 0 0 O 0 0 0 0 0 0 0 0 0 0 4 .50 4 .48 l .13 7 .88 6 .73 9 1.06 6 .73 17 2.06 11 1.2 0 0 0 0 1 .10 1 0 0 .36 4 .42 5 0 0 0 0 76 54 APPENDIX F D f__ hr 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 .06 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 .09 #000000000000HO0HOH000000 000000000000 .09 Total # N Tot # D Grand hr D N NOHO§0000000 I—N .— 0500‘ 00MVHHI—HH 109 0000000 .50 .73 .13 1.12 .85 1.88 1.09 2.74 1.78 .11 .10 .10 .10 .09 .62 .42 [900000000000—0HO0I-‘0I-t000000 hr Total 0 0 0 0 0 0 0 0 0 0 0 0 .06 1 0 4 .06 7 0 l 0 9 .06 8 0 16 .06 10 0 24 0 16 0 l 0 1 0 1 0 1 0 1 0 7 0 5 0 0 .18 2 115 55 NIGHT/ DAY SAMPLE DATA g. coracinus N D _ #__ L #__ L Total _f_N_ Tot _#_D Grand hr hr hr hr N hr D hr Total 4-5/May o 0 0 o 0 0 0 0 o 0 0 0 o 11-12/nny 0 0 2 0.23 0 0 0 0 2 0.23 0 0 2 18-19/May 1 0.12 2 0.23 0 0 0 0 3 0.35 0 0 3 25-26/May o 0 0 0 0 0 0 0 0 0 0 0 o l-Z/Jun 0 0 0 0 0 0 o o 0 0 0 o o 9-10/Jun 0 o 2 0.25 0 0 0 0 2 0.25 0 0 2 16-17/Jun 1 0.12 0 0 1 0.06 0 0 1 0.12 1 0.06 2 22-23/Jun 2 0.25 3 0.38 1 0.06 2 0.12 5 0.62 3 0.19 8 29-30/Jun 2 0.24 1 0.12 3 0.19 0 o 3 0.36 3 0.19 6 6-7/Jul 2 0.26 0 o 0 0 0 0 2 0.26 0 0 2 13-14/Ju1 1 0.12 4 0.50 1 0.06 1 0.06 5 0.62 2 0.12 7 20-21/161 3 0.36 4 0.48 0 0 1 0.06 7 0.85 1 0.06 8 27-28/Ju1 4 0.47 3 0.35 1 0.06 0 0 7 0.92 1 0.06 8 3-4/Aug 2 0.24 5 0.61 3 0.13 0 0 7 0.85 2 0.13 9 10-11/Aug 0 0 1 0.11 0 0 1 0.07 1 0.11 1 0.07 2 17-18/Aug 3 0.33 7 0.78 0 0 1 0.07 10 1.11 1 0.07 11 24-25/163 0 0 3 0.32 0 0 0 0 3 0.32 0 0 3 3l-Aug/1-Sep 1 0.10 2 0.20 0 o 0 0 3 0.31 0 0 3 7-8/Sep 0 0 o 0 0 o 0 0 o o 0 0 0 13-14/Sep 0 0 o o 0 0 0 0 0 0 0 o 0 21-22/s6p 0 0 0 0 0 0 0 0 0 0 0 0 0 28-29/Sep 0 0 0 0 0 0 0 0 0 0 0 0 0 5-6/0.: 0 0 0 o 0 0 0 0 0 o 0 0 o 12-13/0ct 0 o o . 0 0 0 0 0 o 0 0‘ 0 0 26-27/0ct 0 0 0 0 o 0 1 0.09 o 1 1 0.09 1 TOTALS 22 10 7 61 17 NIGHT/DAY SAMPLE DATA g. melanarius 4-5/MBY ll-lZ/May 18-19/Msy 25-26/May l-Z/Jun 9-10/Jun 16-17/Jun 22-23/Jun 29-30/Jun 6-7/Jul 13-14/Ju1 20-21/Jul 27-28/Ju1 3-4/Aug 10-ll/Ang 17-18/Aug 24-25/Aug 3l-Aug/1-Sep 7-8/Sep 13-14/Sep 21-22/Sep 28-29/Sep 5-6/0ct 12-13/0ct 26-27/0ct TOTALS OOQI—oHNHHNomer—wr—NHOOONNO N \O L hr 0.23 0.23 0.12 0.25 0.12 0.39 0.12 0.61 0.12 0.36 0.22 0.11 0.10 0.20 0.09 0.09 HHOOOOOHI—O‘HHOGNwI—OQOHHI—OO N 0 I'I: hr 0.12 0.12 0.12 0.12 0.39 0.25 0.73 0.12 0.11 0.67 0.11 0.10 000000 56 000000000NUIF‘WF‘00000F‘00000 g—I w SI“ 0000900000 0 O3 0 0.06 0.19 0.06 0.33 0.13 000000000 0"00°0F‘0NN5‘0U1—N00000000000 p... 0 hr 0000000000 0 0.12 0.06 0.19 0.2 0.27 0.14 0.14 0.07 0.09 N pa FwONNHOHHwNO OHOHOHNNNmHéI-o U! UI hr 0.23 0.35 0.12 0.12 0.12 0.25 0.24 0.77 0.37 1.33 0.12 0.48 0.11 0.89 0.22 0.20 0.20 0.09 0.09 0.08 D H0000F‘0NNO‘0##U00000F‘00000 U N 0000900000 0 O‘ 0 0.19 0.25 0.25 0.53 0.40 0.13 0.14 0.07 0.09 Total # N Tot # D Grand hr Total UGNNHI—IHP‘WNO p-I 0Uk 1—0 b0 HHOHONNfib 0 00 NIGHT/DAY SAMPLE DATA S. impunctatus 4-5/May 11-12/May 18-19/May 25-26/May l-2/Jun 9-10/Jun 16-17/Jun 22-23/Jun 29-30/Jun 6-7/Ju1 13-14/Jn1 20921/Ju1 27-28/Jul 3-4/Aug lO-ll/Ang 17-18/Aug 24-25/Aug 31-Ang/1-Sep 7-8/Sep 13-14/Sep 21-22/Sep 28-29/Sep 5-6/Oct 12-13/ Oct 26-27/0ct TOTALS 000000000N0—0UIUI3‘N0000000000 p—n \0 gr 000000000 0 0.25 0.48 0.59 0.61 0.11 0.22 0 00000000 N000000000 10 3-0 N 10 000000000U‘NN U! 0‘ w 0000000 0.36 0.26 1.25 2.06 1.18 0.85 0.23 0.56 0 00000000 57 o0o0000r-‘00N0000—09—00000000 El“ 00000000000000 0.13 0.07 000000 00000000NNHNI—0NH00000000.0 g... ‘0 hr 00000000 0 0.06 0.12 0.5 0.06 0.13 0.07 0.13 0.14 0000000 N N000000000 12 l-‘l-‘N NUIFi 000000000NN N UI hr 00000000 0.36 0.26 1.50 2.54 1.76 1.45 0.34 0.78 0 00000000 D 0°00000FNNle—GUHH00000000 N .fi hr 0000000 0.06 0.06 0.19 0.50 0.06 0.13 0.20 0.13 0.13 0.07 000000 Total # N Tot # D Grand Total 0&00000000 r—I—Nr- 900$)! 0000000F‘N‘00‘ 0 \D 58 APPENDIX F 24 Hour Trap Catches g. ingratus g. coracinus g. melanarius Total Total Total 4-5/Mey 0 0 O 0 0 0 0 0 0 11-12/Msy 0 O O 0 0 O 0 0 0 18-19/May 0 0 0 l 1 2 0 0 0 25-26/May 0 0 0 O 0 0 0 0 0 1-2/Jun 0 0 0 0 3 3 0 0 0 9-10/Jun 0 0 0 1 0 1 0 0 0 16-17/Jun 0 0 O 0 1 1 0 0 0 22-23/Jun 1 3 4 2 1 3 0 0 0 29-30/Jun 5 1 6 5 3 8 0 2 2 6-7/Ju1 7 1 8 1 2 3 0 2 2 13-14/Ju1 1o 1 11 3 3 6 0 2 2 20-21/Ju1 5 2 7 0 4 4 0 3 3 27-28/Ju1 1 2 2 4 O 0 0 3-4/Aug 3 0 4 4 O 3 3 10-11/Aug 8 4 12 2 1 3 1 1 2 17-18/Aug 11 1 12 1 3 4 1 5 6 24-25/Aug 3 l 4 3 3 6 0 2 2 31-Aug/1-Sep 1 1 2 1 0 1 0 0 0 7-8/Sep O 0 0 0 0 0 O 0 0 13-14/Sep 1 1 2 0 O 0 0 0 0 21-22/Sep 0 1 1 0 0 0 O 0 0 28-29/Sep 6 3 9 0 0 0 0 0 0 5-6/Oct 1 1 2 O 0 O O 0 0 12-13/Oct 1 0 1 O 0 0 0 0 0 26-27/Oct 2 2 4 0 0 0 0 0 0 TOTALS 64 27 91 22 31 53 2 20 22 4-5/May 11-12/May 18-19/May 25-26/May 1-2/Jun 9-10/Jun 16-17/Jun 22-23/Jun 29-30/Jun 6-7/Ju1 13-14/Ju1 20-21/Ju1 27-28/Ju1 3-4/Aug 10-11/Ang 17-18/Aug 24-25/Aug 31-Aug/1-Sep 7-8/SeP 13-14/Sep 21-22/Sep 28-29/Sep 5-6/Oct 12-13/0ct 26-27/0ct TOTALS _P_. pensylvanicus 0 0 17 11 5 4 3 21 16 10 10 9 23 12 14 4 31 10 15 2 0 0 1 0 0 1 2 0 1 1 0 1 4 6 6 3 3 2 7 14 1 4 1 3 190 116 59 24 Hour Trap Catches Total 28 12 37 20 28 19 18 41 17 HNNHHO 10 21 306 S3 impunctatus I—NNN H0§O~ 000000000305 115 fibrin—0000000 b0NI-90000000 .— ét—o 1 000000H00000 VI 0 Total NNN0000000 HHH41>ww NNO0U|0 0000000-‘009 173 60 LITERATURE CITED Adis, J. 1979. Problems of interpreting arthropod sampling with pitfall traps. Zool. Anz., 202:177-184. Andrews, A. W. 1921. The Coleoptera of the Shires expedition to whitefish Point, Chippewa County, Michigan. Mi. Acad. Sci. Arts, Lets, Papers, 13293-3900 Basra, M. A., 1979. Patterns of movement of radioactive Carabid Beetles. Oecologia, 44:125-140. Barlow, C. A. 1970. Phenology and distribution of some Pterostichus of Eastern Canada. J. N.Y. Ent. Soc., 78:215-236. Bauer, Thomas. 1977. 'The relevance of the brightness to visual acuity, predation, and activity of visually hunting ground-beetles (Coleoptera, Carabidae). Oecologia, 30:63-73. Beck, 8. D. 1980. Insect Photoperiodism, 2nd Ed. Academic Press, Toronto. 387 pp. Boiteau, G. 1983. Activity and distribution of Carabidae, Arachnida, and Staphylinidae in New Brunswick potato field's Can. Ent., 115:1023‘10300 Boucher, J. F. and J. C. Malauss. 1984. 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