USDA unitedStates AgricultureDepartment of and Carabid Beetle l:_,=c_ Species Associated with _:::S::t,a'"_0 Ecological Landtype Station , GeneralTechnical Phases in

Report NC-201 Northern Michigan

Timothy T. Work, DeborahG. McCullough, and William J. Mattson

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North Central Research Station Forest Service--U.S. Department of Agriculture 1992 Folwell Avenue St. Paul, Minnesota 55108 Manuscript approved for publication September 14, 1998 1998 Moth and Carabid Beetle Species Associated with Two Ecological Landtype Phases in Northern Michigan

. Timothy T. Work, Deborah G. McCullough, and William J. Mattson

Insects are among the truly critical components test the hypothesis Ho: faunal communities of forest ecosystems, affecting many key pro- are uniquely asembled for each unique forest cesses such as decomposition and nutrient ecosystem, and in so doing, we also provide a cyc!ing, regulation of primary productivity, and faunal inventory of two major groups for pollination. Moreover, they are typically the two common forest ecosystem units in the backbone of most food webs, serving both as Huron-Manistee National Forest in the north- prey--the primary protein and mineral sources ern lower peninsula of Michigan. for a wide variety of , and as important, influential predators and parasitoids them- Ecological classification systems (ECS) provide selves, helping" to regulate populations of an integrated framework for studying ecosys- potentially damaging herbivores, tems at multiple spatial and temporal scales (Sims et a/. 1996). In many national forests in Since ecosystem management has become an the United States, these hierarchical systems accepted perspective nearly worldwide in are used to identify ecological capabilities or sustainable resource management (Brooks and constraints associated with specific areas or Grant 1992, Christensen et al. 1996, Kessler et sites. An ECS can also be used as a basis for a/. 1992, Thomas 1996), those developing and monitoring changes over time. An ECS re- _nplementing such complex management plans cently developed for the Huron-Manistee must have broader and deeper knowledge National Forest in northern lower Michigan has about all facets of ecosystems (Brooks and as its fundamental unit the ecological landtype Grant 1992, Christensen eta/. 1996). Not only phase (ELTP) (Cleland et al. 1994). ELTP's is this knowledge necessary to initially set have distinct softs, landscape positions, and sound ecosystem management goals, but it is natural vegetative communities. ELTP's are also important to assess whether management sub-units of ecological landtypes (ELT's), which goals are being attained. Although the impor- in turn are sub-units of landtype associations tance of in forests is widely recognized, (LTA's). LTA's often encompass thousands of Very few studies have attempted to describe acres and are defined by major geomorphology particular insect communities associated with and vegetation features. specifi c forest ecosystems. In this paper, we . Plant species composition and structure of overstory and understory vegetation typically have a major influence on diversity, abun- Timothy T. Work is a Research Assistant, dance, and speciescomposition of insect Department of Entomology, Michigan State communities {Strong et aL 1984}. We hypoth- University,East Imnsing,MI (currentlywith the esized,therefore,that insectcommunities are Department of Entomology, Oregon State likelyto differbetween ELTP's, even though University,Corvallis,OR); Deborah G. they may be spatiallyproximate to each other. McCullough isan Associate Professor,Depart- Descriptionof insectcommunities associated ment of Entomology and Forestry,Michigan with specificELTP's would provide resource StateUniversity,East Lansing,MI; and Will- managers with valuable biologicalinventory Jam J. Mattson isa Research Entomologist, informationand limitthe need to repeatedly Nort_ Central Research Station,Rhinelander, conduct time- and labor-intensivesurveys. WI. We surveyed two major groups of insects in two National Forest. On each site, tree species hdrdwood ELTP's in the Huron-Manistee composition and basal area were recorded in National Forest during the summers of 1993, five variable radius plots using a BAF 10 prism. 1994, and 1995. In this publication, we pro- vide a list of the species of (mainly Although gypsy moth populations were expand- ) and carabid beeries that were associ- ing through much of northern Michigan in the ated with these two ELTP's. Both insect groups late 1980's and early 1990's, no visible defolia- were sampled in four replicate sites classified tion occurred in the Manistee portion of the as ELTP 20 and in four sites classified as ELTP Huron-Manistee National Forest until 1993 (D. 45. ELTP 20 sites are dominated by northern McCullough, unpubl, data; S. Katovich, USDA red oak (Quercus rubra L.), white oak (Q. a/ha Forest Service, NA State & Private Forestry, L.) and a low-diversity ground flora (Cleland et pers. comm.). From 1993 to 1995, gypsy moth al. 1994). Soils are typically sandy and xeric populations became established in localized with medium to low productivity (Host et a£ areas of the Manistee, causing moderate to 1987, 1988). In contrast, ELTP 45 sites are high defoliation in some stands. Outbreak- dominated by sugar maple (Acer saccharum level populations of gypsy moth typically Marsh), with a minor component of American persisted for 1 to 2 years, then subsided. We basswood (Tilia americana L.) (Cleland et a£ estimated the size of gypsy moth populations 1994): Other less abundant overstory species, using mean egg mass counts from four 0.01-ha including.red oak, quaking aspen (Populus plots located in each study site (Kolodny-Hirsch .tc:emuloides Michx.), black cherry (Pn_us 1986). Defoliation of overstory trees was serotina Ehrh.), and white ash (Frax/nus visually estimated in each stand as high (75 to amer/cana L.), are often present. ELTP 45 sites 100 percent), moderate (50 to 75 percent), low also have a notably rich herbaceous under- (25 to 50 percent), or negligible (<25 percent). Story, well-drained mesic soils, and high pro- Defoliation was further quantified in 1994 and ductivity (Host et a/. 1987, 1988; Zak and 1995 with a ceptometer (Sunfleck TM )I. Sensors Pregitzer 1990; Zak et a£ 1986, 1989). along the ceptometer quantified photosyntheti- cally active radiation (PAR) penetrating the Lepidoptera and carabid ground beeries are canopy, and readings were used to estimate diverse taxa that are important in various leaf area index (LAI) in defoliated and forest ecosystem processes. More than 500 undefoliated stands (Work 1996). . species of macrolepidoptera (M. Nielsen, MSU Dept. of Entomology, pers. comm.) and 38 EI,TP 20 Stands species of ground beeries (Liebherr and Mahar 1979) have been recorded in northem lower Two oak-dominated ELTP 20 stands were Michigan. Defoliating Lepidoptera can affect located at 44 ° 00' N, 86 ° 00' W, near the town of nutrient cycling, tree mortality, and growth Branch, and two other stands were located at rates, and can alter stand structure (Mattson 44 ° 08' N, 86 ° 09'W, near the village of Freesoil, and Addy 1975). Many carabids are important all in Mason County. Although northern red predators of other including notori- oak and white oak dominated the overstory, big ous pest species such as gypsy moth tooth aspen (Populus grandidentata Michx.) and (Lymantria dispar L.) (Cameron and Reeves red maPle (Acer rubrum L.) together made up 1990, Weseloh et a£ 1995). Both lepidopteran close to 30 percent of the canopy in all stands. and Carabid species may be sensitive indicators Overstory trees were approximately 20 m tail. of Habitat quality or disturbance (Martel and A sparse understory of red maple saplings and Mauffette 1997, Niemela et a£ 1993). witch hazel (Hamamelis virginiana L.) of less than 3 cm diameter breast height (d.b.h.) was METHODS also present.

Study Area Gypsy moth populations caused severe defolia- tion at Branch sites in 1993 and moderate to Lepidoptera and carabid beeries were sampled severe defoliation at Freesoil sites in 1994 and from eight stands in the Huron-Manistee 1995 (table 1). Heavily defoliated stands had National Forest in northern lower Michigan. We randomly chose four stands classified as • ELTP 20 and four stands classified as ELTP 45 1Mention of trade names does not constiage from maps provided by the Huron-Manistee endoresement by the USDA Forest Service. 2 Table l.---Mean density (+_SE) of gypsy moth egg masses per hectare and defoliation estimates in "ELTP 20 and ELTP 45 stands in northern lower Michigan during the summers of 1993 through 1995

ELTP Site 1993 1994 1995

20 Branch

.. No. egg masses 1,293 __.119.2 849 +_302.5 49 _ 17.3 Defoliation High None None

20 Freesoil No. eggmasses 5 _+2.5 233 __27.1 1942 __+388.0 Defoliation None Moderate High

45 Harrietta No. eggmasses 45 _ 17.9 133 +_29.0 52 +_1_7.4 Defoliation None None None

45 Mesick No. eggmasses 187 +_17.9 168 _+42.5 11+__3.8 Defoliation None None None

roughly 60 percent of the LAI of undefoliated 15, and August 10-12 from ELTP 20 sites and stands (Work 1996). Trees in stands that had on May 21-22, June 22-23, July 20-22, and been heavily defoliated re-flushed by late July, August 18-20 from ELTP 45 sites. In 1994, and no tree mortality was observed during the insects from ELTP 20 sites were collected on course of this 3-year study. May 10-12, June 15-16, July 11-13, and August 9-10 and from ELTP 45 sites on May EL3r_P45 Stands 18-20, June 20-21, July 19-20, and August 15- 17. In 1995, collection dates were May 15-17, Two northern hardwood ELTP 45 stands were June 12-14, July 10-11, and August 15-16 on located near the town of Mesick at 44 ° 22' N, ELTP 20 sites and May 22-24, June 19-20, 86 ° 44' W, and two similar sites were located July 17-19, and August 21-22 on ELTP 45 near the village of Harrietta at 44 ° 19' N, 86 ° 44' sites. Sampling was delayed until the following W, all in Wexford County. Northern hardwood day if rain or high winds occurred.. species including sugar maple and American basswood dominated the overstory, although Canopy Stratum other species including black cherry and beech (Fagus grandifolia Ehrh.) were scattered on the In 1993, the overstory canopy was sampled Sites. Overstory trees were approximately 30 m using pole pruners, modified sweepnets with tall, and sites had a dense understory of l-m- 10-m extending handles, and suspendable UV tall sUgar maples of less than 3 cm. Gypsy traps. In 1994 and 1995, pole pruners and moth populations were negligible in ELTP 45 extended sweepnets were not used because of sites because few preferred host species were the relatively low number of specimens col- present (Liebhold eta/. 1995). lected relative to the sampling effort needed.

Insect Sampling Pole pruners were used in 1993 to clip branch tips, 1 m in length, at 20-m intervals along a Lepidoptera and carabids were collected from 120-m transect located randomly through the three stratamthe overstory canopy, the shrub stand. Clipped branches were placed in plastic stratum, and the groundDin each stand, bags until they could be examined for insects. Insects were collected four times during the In 1993, sweepnets with extending handles summer in 1993, 1994, and 1995 at approxi- were used to sweep lower canopy foliage at 20- • mat_ly 4-week intervals. In 1993, insects were m intervals along a 120-m randomly located collected on May 12-13, June 15-17, July 13- transect through each stand. Four sweeps of 3 the lower canopy foliage with a 50-cm diameter Traps were left open for 24 hours during each " net were made at each stop along the transect, monthly sampling period.

Ultraviolet traps were used in all 3 years, and Processing and Taxonomic Identification each consisted of a 22-watt UV bulb connected to a photosensor and powered by a 6-volt Adult Lepidoptera were placed in plastic con- motorcycle battery. UV traps were operated tainers lined with tissue paper and stored in a approximately 8 hours beginning at dusk freezer until i_entified. Only adult Lepidoptera during each monthly sampling period. A were identified to species because few larvae collection funnel and bucket loaded with were collected and resources did not permit the insecticidal pest strips (Vapona TM) were at- rearing of immatures. All Lepidoptera speci- tached below the bulb. Ropes were suspended mens were pre-sorted and recorded by by Shooting an arrow with an attached line over parataxonomists at Michigan State University, a large limb in the middle to upper canopy of a East Lansing, MI. Species identifications of tree at the center of the stand. UV traps were Lepidoptera were made by John Wilterding, then raised into the lower canopy. Use of a low Robert Kriegel, and Mogens Nielsen at the wattage bulb and a rain cover over the bulb Michigan State University Center for Insect typically restricts potential long-distance Diversity. Carabids were stored in70 percent attraction of nocturnal insects, resulting in a EtOH. Carabid identifications were made by T. localized sample reflecting endemic insect Work using taxonomic keys developed by populations (Bowden 1982). Lindroth (1969); taxonomy followed Bousquet and Larochelle (1993). Voucher specimens for Shrub Stratum all taxa were deposited at the Center for Ar- thropod Study, Department of Entomology, Shrub insects were collected using malaise Michigan State University, East Lansing, MI. • traps and sweepnets. One malaise trap with an insecticidal pest strip was set up in the RESULTS center of each stand and left open for 24 hours during each monthly sampling period. Shrub Lepidoptera vegetation (0.5 to2 m high) was sampled using a 38-cm diameter sweepnet. Sixteen sweeps Over the course of this 3-year project, 12,346 (four from each cardinal direction) of vegetation adult Lepidoptera were collected. The vast . , at 20-m intervals were made along a 120-m majority (94 percent, 11,646 insects) came randomly located transect through the stand from UV traps and 5 percent (673 insects) between 1000 and 1600 hour. came from malaise traps. A total of 489 Lepi- doptera species was collected from the two Ground Stratum ELTP's (table 2], Four families, , Noctuidae, Lyonetiidae, and Geometridae, Ground insects were sampled using 10 accounted for 77 percent of all Lepidoptera unbaited Pitfall traps located at 10-m intervals collected (table 3). Of the Lepidoptera/species, along a randomly selected transect through the 169 were common to both ecosystems, which stand. Pitfall traps had a diameter of 10.5 cm, accounted for 89 percent (10,929 insects) of the contained soapy water, and were covered with a total Lepidoptera collection. Collection dates, four-legged *roOf" to minimize capture of non- location, and total abundance of Lepidoptera 'ground insects and prevent predation by birds, are reported in table 4.

Table 2._Total number of Lepidopteran families, species, and individuals coUected from ELTP 20

and ELTP 45 stands in northern ¢ lower Mich_an during the summers of 1993 through 1995

ELTP 20 ELTP 45 Grand total

Year 1993 1994 1995 Total 1993 1994 1995 Total No. families 25 23 26 37 24 21 25 33 39 No. species 171 133 168 343 152 118 173 315 489 • No. individuals 1,604 1,691 2,183 5,478 1,994 1,140 3,736 6,870 12,348

4 .Table 3.--Five most abundant families of Lepidoptera coUected from ELTP 20 and ELTP 45 Stands in northern lower Michigan during the summers of 1993 through 1995

Family - No. individuals collected Relative abundance in percent • ,

Tortricidae 5,996 49 Noctuidae 1,809 15 Lyonetiidae 917 7 Geometridae 739 6 Others (35 families) 2,887 23

ELTP 20 had 156 species that were not found Twenty-five carabid species were collected in in ELTP 45. Of these unique species, 17 both ELTP's, accounting for 93 percent (2,355 percent (154 individuals) were Hydrimoena sp. beeries) of the total catch. Five species com- (Geometridae) and 6 percent (53 individuals) prised 60 percent of the total catch, namely were Blephanomastix ranalis (Gn.) (Pyralidae). Stenopholus comma (F.), Carabus goryi Dej, The remaining 77 percent included 154 spe- Synuchus impunctatus (Say), Bembidion cies; less than 50 individuals of these species partreule Dey., and Stenopholus ochropezus were collected. Of the 154 species, 87 were (Say) (table 7). TWenty-one species were represented by a single specimen, unique to ELTP 20; nine of these are typically found in xeric, well-lighted habitats. Twelve ELTP 45 had 129 unique species that were not species were unique to ELTP 45; nine of which collected in ELTP 20 stands. ELTP 45 collec- are associated with moist, shady forest habitat. tions were dominated by the two tortricids Specialist predators of mollusks such as Pandemis canadana Kft. (21 percent, 114 Dicaleus teter Bon., Scaphinotes bilobus (Say), individuals) and Proteoteras moffatiana Fern. and Spheroderus stanostanus lecontei Dej. were (15 percent, 83 individuals). Although found only in ELTP 45. Pandemis canadana was collected in high numbers, it was collected only in July 1995, DISCUSSION . and only at one site (table 4). Proteoteras moffatiana, a bud-boring pest of sugar maple, Although many of the abundant was collected in all 3 years and at all four ELTP species were present in both ELTP 20 and 45 sites (table 4). The remaining 64 percent of ELTP 45 stands, the less abundant species the unique Lepidoptera were comprised of 127 may be better indicators of the differences species; 64 of these species were represented between these two ecosystems. Within an • by a single individual, ecosystem, insect abundance and diversity are determined by a combination of factors such as Carabid Beeries food availability (whether plants or prey), food quality, climate, and habitat structure (Strong Over 3 years, a total of 2,528 carabids repre- et al. 1984). Abundant species may be com- senting 59 species was collected (table 5). Most mon across ecosystems because they are less carabids were collected with UV traps (61 tightly coupled with these factors or because percent of total catch; 1,537 individuals) and some of these factors are universally available pitfall .traps (39 percent of total catch, 988 across ecosystems. Uncommon species may individuals). Collection dates, location, and have more stringent requirements, and thus total abundance of, carabid species are reported may be more specific to a single habitat or in table 6. Small brachypterous species (short- ecosystem. In this study, ELTP 20 and ELTP • winged beetles) from the genera Agonurr_ 45 had 156 and 129 unique species of Lepi- Bembidion, Harpalus, and Stenopholus were doptera, respectively. Although these species mostly collected in UV traps. Imrger species do not dominate numerically, they clearly (both apterous and/or poor fliers) in the genera suggest that the insect communities in these Carabus, Myas, and Pterostichus were collected ecosystems do indeed differ. primarily in pitfall traps.

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L • Table 7.mFive most abundant carabid species collected from ELTP 20 and ELTP 45 stands in northern lower Michigan during the summers of 1993 through 1995

Carabid Species No. beetles collected Relative abundance in percent

Stenopholus comma (F.) 522 21 Carabus goryi Dej. 345 14 Synuchus impunctatus (Say) 234 9 _Bembidion versicolor Dej. 214 8 Stenopholus ochropezus (Say) 213 8

Species richness and abundance data for methods can increase the resolution of uncommon species should always be inter- biodiversity inventories without requiring preted carefully, with particular attention to survey of additional sites or greatly increasing sampling intensity and efficiency. When cost and effort of sampling. species are represented by a single individual, ecological interpretations of data are limited to Our results should provide baseline data for presence-absence statements. However, re- managers interested in quantifying, conserving, sults from our survey provide a framework to or increasing insect species diversity in these begin more intensive investigations of host- ecosystems. This information may also be plant associations of rare species that appear useful for entomologists or managers attempt- to be unique to one ELTP. ing to predict effects of natural disturbance or silvicultural activities on forest insect commu- While Lepidoptera communities are probably nities. linked closely to vegetation characteristics, Carabid diversity and abundance may more ACKNOWLEDGMENTS closely reflect microclimatic gradients. In ELTP 20, almost 50 percent of the unique species are We gratefully acknowledge Brian Kopper for his reported to be associated with well-lit, xeric assistance in collecting and processing all of habitats. In contrast, in ELTP 45, 75 percent the specimens. Dr. Fred Stehr and Dr. Robert " of the unique species are considered either Haack reviewed an earlier draft of the manu- hygrophagous (associated with humid environs) script and provided many helpful comments. or associated With prey found in moist habi- Funding for this study was provided by the tats. The difference in carabid fauna may McIntire-Stennis Cooperative Forestry Re- correspond to ecosystem differences in habitat search Program. Additional support was features that affect microclimate. ELTP 20 had provided by the Michigan Agricultural Experi- an open overstory canopy and little understory ment Station Project No. MICLO 1710. and ground vegetation, allowing more sunlight to reach, the forest floor, increase temperature, LITERATURE CITED and decrease relative humidity. On the other hand, ELTP 45 had a dense canopy and rich Bousquet, Y.; Larochelle, A. 1993. Catalogue , understory and ground flora, which presum- of the Geadephaga (Coleoptera: ably led to higher relative humidity and cooler, Trachypachidae, Rhysodidae, Carabidae moister conditions, includng Cicinidelini) of America north of Mexico. Memoirs of the Entomological The majority of both Lepidoptera and carabids Society of Canada. 397: 1-111. were Collected in l_ traps. This trapping • method, while biased, provided many speci- Bowden, J. 1982. An analysis of factors mens with relatively little effort and cost. affecting catches of insects in light- Carabid beeries are commonly sampled using traps. Bulletin of Entomological Research. pitfall traps because of the ease and low cost of 72: 535-556. maintaining traps. However, our results showed a Significant proportion of the carabid Brooks, D.J.; Grant, G.E 1992. New perspec- • fauna (61 percent) was underrepresented by tives in forest management: background, pitfalls only. The use of multiple trapping science issues, and research agenda. Res. 23 . Pap. PNW-456. Portland, OR: U.S. Depart- Liebherr, J.; Mahar, J. 1979. The carabid ment of Agriculture, Forest Service, Pacific fauna of the upland oak forest in Michi- Northwest Research Station. 17 p. gan: survey and analysis. Coleopterists Bulletin. 33:183-197. Cameron, E.A.; Reeves, R.M. 1990. Coleoptera associated with gypsy moth, Lymantria Liebhold, A.M." Gottschalk, K.W." Muzika, R.M." dispar (L.) (Lepidoptera:Lymantriidae), Montgomery, M.E." Young, R.; O'Day, K.; populations subjected to Bacillus Kelly, B. 1995. Suitab!i!!ty of North thuringiensis Berliner treatments in American tree species to gypsy moth: a Pennsylvania. Canadian Entomologist. summary of field and laboratory tests. 122: 123-129. Gen. Tech. Rep. NE-211. Radnor, PA: U.S. Department of Agriculture, Forest Service, Christensen, N.L.; Bartuska, A.M.; Brown, Northeastern Research Station. 34 p. J.H.; Carpenter, S.; .D'Antonio, C.; Francis, R.;Franklin, J.F.; MacMahon, J.A.; Noss, Lindroth, C.H. 1969. The ground-beetles of R.F.; Parsons, D.J.; Peterson, C.H.; Turner, Canada and Alaska. Opuscula M.G.; Woodmansee, R.G. 1996. The report Entomologica Supplenta. 20, 24, 26, 29, of the Ecological Society of America 33, 34, 35: I-1192. committee on the scientific basis for ecosystem management. Ecological Martel, J.; Mauffette, Y. 1997. Lepidopteran Applications. 6:665-69 I. communities in temperate deciduous forests affected by forest decline. Oikos. Cleland, D.T.; Hart, J.B.; Host, G.E.; Pregitzer, 78: 48-56. K.S.; Ramm, C.W. 1994. Field guide" ecological classification and inventory Mattson, W.J.; Addy, N.D. 1975. Phytopha- system of the Huron-Manistee National gous insects as regulators of forest Forests. Cadillac, MI: U.S. Department of primary production. Science. 190:515- Agriculture, Forest Service, Huron-Manistee 522. National Forest. [varied pages]. Niemela, J.; Langor, D.; Spence, J.R. 1993. Host, G.E.; Pregitzer, K.S.; Ramm, C.W.; Hart, Effects of clear-cut harvesting on boreal J.B.; Cleland, D.T. 1987. Landform-medi- ground-beetle assemblages (Coleoptera: ated differences in successional path- Carabidae) in Western Canada. Conserva- ways among upland forest ecosystems in tion Biology. 7:551-561. northwestern lower Michigan. Forest Science. 33: 445-457. Sims, R.A.; Corns, I.G.W.; Klinka, K. 1996. Global to local: ecological land classifica-

• ." Host, G.E.; Pregitzer, K.S.; Ramm, C.W.; Lusch, tion. In: Proceedings of the international D.P." Cleland, D.T. 1988. Variation in conference; Thunder Bay, Ontario, Canada; overstory biomass among glacial land- 14-17 August 1994. Environmental Moni- forms and ecological land units in north- toring and Assessment. 39: 1-3. western Lower Michigan. Canadian Journal of Forest Research. 18: 659-668. Strong, D.A.; Lawton, J.H.; Southwood, T.R.E. • 1984. Insects on plants: community Kessler, W.B.; Swasser, H.; Cartwright, C.W.; patterns and mechanisms. Cambridge, Caplan, J.A. 1992. New perspectives for MA: Harvard University Press. 313 p. sustainable natural resource manage- ment. Ecological Applications. 2:221-225. Thomas, J.W. 1996. Forest Service perspec- tive on ecosystem management. Ecologi- Koldony-Hirsh, D.M. 1986. Evaluation of cal Applications. 6: 703-705. methods for sampling gypsy moth (Lepi- doptera: Lymantriidae) egg mass popula- Weseloh, R.; Bernon, G.; Butler, L.; Fuester, R.; tions and development of sequential McCuUough, D.; Stehr, F. 1995. Releases sampling plan. Environmental Entomology. of Calasoma sycophanta (Coleoptera: 15: 122-127. Carabidae) near the edge of gypsy moth 0Lepidoptera: Lymantriidae) distribution. Environmental Entomology. 24:1713-1717. 24 Work, T.T. 1996. Impacts of gypsy moth (Lepidoptera: Lymantriidae) on native arthropod abundance, species richness, and diversity in two hardwood ecosys- tems in northern lower Michigan. East Lansing, MI: Michigan State University, Department of Entomology. 196 p. M.S. thesis.

Zak, D.R." Pregitzer, K.S. 1990. Spatial and temporal variability of nitrogen cycling in northern lower Michigan. Forest Sci- ence 39" 367-380.

Zak, D.R." Host, G.E." Pregitzer, K.S. 1989. Regional variability in nitrogen mineral- ization, nitrification, and overstory biomass in northern Lower Michigan. Canadian Journal of Forest. Research. 19" 152i-1526.

Zak, D.R.; Pregitzer, K.S.; Host, G.E. 1986. Landscape variation in nitrogen mineral- ization and nitrification. Canadian Jour- nal of Forest Research. 16:1258-1263.

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_v.s. 6OVERNM_PrRINTINGOmC_ t_. 7_mvsm_s 25 Work, Timothy T." McCullough, Deborah G." Mattson, William J. 1998. Moth and carabid beetle species associated with two eco- logical landtype phases in northern Michigan. Gen. Tech. Rep. NC-201. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Research Station. 25 p.

More than 12,300 moths and 2,500 carabid beetles were trapped during 3 years (1993 through 1995) in two different : ecological land type phases (ELTP's) in the Huron-Manistee National Forest in Michigan. One ELTP (no. 20) was dominated by oaks, and the other (no. 45) was dominated by sugar maple; each had distinctive kinds of insects, in spite of the fact that many species were shared. Collection dates, trapping method, and the total abundances of each of the 489 moth species, and each of the 59 carabid species are reported.

Key Words: biodiversity, ecosystem management, ELTP, LTA, faunal list.

o Our job at the North Central Forest Experiment Station is discovering and creating new knowledge and technology in the field of natural resources and conveying this information to the people who can use it. As a new generation of forests emerges in our region, managers are confronted with two unique challenges: (1) Dealing with the great diversity in composition, quality, and ownership of the forests, and (2) Reconciling the conflicting demands of the people who use them. Helping the forest manager meet these challenges while protecting the environment is what research at North Central is all about.

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