Great Lakes Entomologist the GREAT LAKES ENTOMOLOGIST Published by the Michigan Entomological Society Vol

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The Great Lakes Entomologist THE GREAT LAKES ENTOMOLOGIST Published by the Michigan Entomological Society Vol. 46, Nos. 1 & 2 Spring/Summer 2013

Volume 46 Nos. 1 & 2 ISSN 0090-0222

Table of Contents THE

Variation in Lepidopteran Occurrence in Hemlock-Dominated and Deciduous-Dominated Forests of Central Appalachia GREAT LAKES L. E. Dodd, Z. Cornett, A. Smith, L. K. Rieske...... 1 Attraction of Agrilus planipennis (Coleoptera: Buprestidae) and Other Buprestids to Sticky Traps of Various Colors and Shapes ENTOMOLOGIST Toby R. Petrice, Robert A. Haack, and Therese M. Poland...... 13 Discontinuity in the Insect Assemblages of a Northern Lower Michigan Stream David C. Houghton, Constance M. Brandin, Leila Reynolds, and Lindsey L. Elzinga...... 31 Diversity and Community Structure of Stream Insects in a Minimally Disturbed Forested Watershed in Southern Illinois J. E. McPherson, Jacqueline M. Turner, and Matt R. Whiles...... 42 Impact of Mecinus janthinus (Coleoptera: Curculionidae) on the Growth and Reproduction of Linaria dalmatica (Scrophulariaceae) Elizabeth J. Goulet, Jennifer Thaler, Antonio Ditommaso, Mark Schwarzländer and Elson J. Shields...... 90 First Report of the Adventive Species Sitochroa palealis (Lepidoptera: Crambidae) in Pennsylvania and its Attraction to the Sex Pheromone of the European Corn Borer,

Ostrinia nubilalis (Lepidoptera: Crambidae) Vol. 46, Nos. 1 & 2 Neelendra K Joshi, David J. Biddinger, Shelby Fleischer, Steven Passoa...... 99 Inventory of Aquatic and Semiaquatic Coleoptera from the Grand Portage Indian Reservation, Cook County, Minnesota David B. MacLean...... 104 Epizooic Diatoms on the Cerci of Ephemeroptera (Caenidae) Naiads Daniel E. Wujek...... 116 Distribution records for Zenoa picea (Palisot de Beauvois, 1805) (Coleoptera: Callirhipidae) from the United States. Edwin L. Freese...... 120 The Horsehair Worm Gordionus violaceus (Nematomorpha: Gordiida) in Minnesota Philip A. Cochran, Jacob D. Zanon, Ben Hanelt...... 133 First Record of Chrysomya rufifacies (Diptera: Calliphoridae) in Wisconsin Jordan D. Marché II...... 135 Occurrence of Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae) Feeding on Cirsium pitcheri Flowers Jordan M. Marshall...... 138 Demonstration of Sex Pheromones in Anabolia bimaculata, Hydatophylax argus, and Nemotaulius hostilis (Trichoptera: Limnephilidae) PUBLISHED BY David C. Houghton...... 142 THE MICHIGAN Cover photo Hydatophylax argus (Trichoptera: Limnephilidae) male-female in copula. ENTOMOLOGICAL Photo by David Houghton 2013 SOCIETY THE MICHIGAN ENTOMOLOGICAL SOCIETY INSTRUCTIONS FOR AUTHORS SUBJECTS 2012-2013 OFFICERS Papers dealing with any aspect of entomology will be considered for publication in The Great Lakes Entomolo- President David Houghton gist. Appropriate subjects are those of interest to professional and amateur entomologists in the North Central President Elect Martin Andree States and Canada, as well as general papers and revisions directed to a larger audience while retaining an Immediate Past President Toby Petrice interest to readers in our geographic area. Treasurer Tina Ciaramitaro All manuscripts are refereed by at least two reviewers. 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OTHER BUSINESS Other correspondence should be directed to the Secretary, Michigan Entomological Society, 2104 Needham Rd., Ann Arbor, MI 48104. 2013 THE GREAT LAKES ENTOMOLOGIST 1

Variation in Lepidopteran Occurrence in Hemlock-Dominated and Deciduous-Dominated Forests of Central Appalachia L. E. Dodd1,2,*, Z. Cornett1, A. Smith3, and L. K. Rieske1

Abstract Eastern hemlock, (Tsuga canadensis Carrière, Pinaceae), is threatened with extirpation by an exotic invasive herbivore, the hemlock woolly adelgid, (Adelges tsugae Annand, Homoptera: Adelgidae). Given this threat, a broader and more detailed knowledge of the community associated with eastern hemlock is merited. As Lepidoptera are important members of forest communities, this study was initiated to determine the relative occurrence of Lepidoptera in hemlock-dominated and deciduous-dominated habitats by evaluating abundance, species richness, temporal variation, and composition overlap. Lepidoptera were surveyed using blacklight traps from May – August 2010 at two collection sites in the Appalachian region of eastern Kentucky. The first collection site was within a forest stand dominated by mixed deciduous species, the second site possessed an overstory of eastern hemlock. Lepidoptera ≥ 20 mm in wingspan were identified and enumerated, yielding a total of 1,020 individuals of ≥ 137 species and 18 families. The total number of Lepidoptera captured in May and June was fewer than in July and August (P ≤ 0.05). The composition of the assemblage varied between collection sites as well as seasonally; 85 species were identified at the deciduous site and 107 species were identified at the hemlock site. While 27 species were recorded only at the deciduous site, 49 species were unique to the hemlock site. Of those unique to the hemlock site, five species were either detritivores or conifer specialists. These data demonstrate the importance of both deciduous and hemlock-dominated forest habitats for many species of Lepidoptera in Appalachia. Our study forms a foundation for understanding species richness patterns of Lepidoptera in hemlock forests in North America and is a useful baseline for comparisons of richness and diversity post invasion by the hemlock woolly adelgid.

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Lepidoptera are among the most abundant and conspicuous of forest insects. Defoliation by these insects is widespread in forests, with pestiferous species found throughout eastern North America (Covell 2005, Summerville and Crist 2008). Beyond the importance of these insects as forest herbivores, Lepidoptera also play a valuable role in the larger food web by serving as critical prey for both vertebrate and invertebrate predators (Dix et al. 1995, Lacki et al. 2007). These insects are found broadly across eastern North America and in mountain regions deciduous forests harbor high species diversity across a gradient of conditions (Dodd et al. 2008, Summerville and Crist 2008, Dodd et al. 2012). While species inventories for this group exist across North America’s eastern deciduous forests (Summerville et al. 1999, Summerville and Crist 2001, Dodd et al. 2011) and northern boreal forests (Thomas and Thomas 1994, Thomas 2001, Pohl et al. 2004), specific knowledge of the Lepidoptera affiliated with forests dominated by eastern hemlock, (Tsuga canadensis Carrière, Pinaceae), is lacking. Eastern

1Department of Entomology, University of Kentucky, Lexington, KY 40546 USA. 2Department of Forestry, University of Kentucky, Lexington, KY 40546 USA. 3531 Whippoorwill Drive, Morehead, KY 40351 USA. *Corresponding author: (e-mail: [email protected]). 2 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 hemlock is a slow-growing, shade tolerant tree that serves as a foundation species in forests of eastern North America (Godman and Lancaster 1990). In the northern portions of its range eastern hemlock grows in large contiguous tracts, but in the south, including central and southern Appalachia, it is most often confined to riparian zones and moist cove sites (Godman and Lancaster 1990). Hemlock is often the only conifer present where it occurs, and it provides critical nesting habitat for birds (Tingley et al. 2002). The canopy is dense, regulating air and soil temperatures (Adams and Loucks 1971) and limiting light penetra- tion to the forest floor. As a riparian tree it influences headwater stream habitat and benthic macroinvertebrate communities (Snyder et al. 2002) and supports unique fish communities (Ross et al. 2003). Hemlock litter decomposes slowly, and associated soils are poor in nutrients (Yorks et al. 2003), acidic, and with a characteristic species-poor understory community (Godman and Lancaster 1990). As a foundation species, eastern hemlock influences the composition and structure of arthropod communities (Snyder et al. 2002, Dilling et al. 2007, Rohr et al. 2009, Mallis and Rieske 2011). In eastern North America eastern hemlock and its community associates are threatened with extirpation by an exotic invasive herbivore, the hemlock woolly adelgid, (Adelges tsugae Annand, Homoptera: Adelgidae). The adelgid is a xylem-feeding insect of Asian origin that depletes the starch reserves of its host plant. Eastern hemlock is highly susceptible, and is rapidly being colo- nized by the adelgid throughout its range. Ecosystems dominated by eastern hemlock are a critical, though uncommon component of the forest landscape throughout much of the central and southern Appalachians. Replacement of eastern hemlock by deciduous species will likely change nutrient cycling and the physical and chemical conditions of terrestrial habitats (Yorks et al. 2000). Given the threat posed to eastern hemlock and associated communities from adelgid-induced hemlock mortality, a broader and more in-depth knowledge of hemlock community associates is merited and timely (Buck et al. 2005, Dilling et al. 2007). To address this, the main objective of this study was to document the relative occurrence of Lepidoptera by determining abundance, species richness, temporal variation, and composition overlap in a hemlock-dominated and deciduous-dominated habitat.

Materials and Methods Surveys were conducted at two sampling sites ca. 300 m apart in a contiguous forest tract in Bath County, Kentucky. These sites lay in the Cumber- land District of the Daniel Boone National Forest at the juncture of Bath and Menifee counties, which is part of the Western Allegheny Plateau (Level III Ecoregion) and include portions of the Knobs-Lower Scioto Dissected Plateau and the Northern Forested Plateau Escarpment (Level IV Ecoregions) (Woods et al. 2002). The overstory of one site was dominated by eastern hemlock and the other was dominated by deciduous species (Acer, Carya, and Quercus spp.). The hemlock site was found at 38°1’1.26” N, 83°35’25.50” W; the deciduous site was found at 38°1’2.35” N, 83°35’15.25” W. Canopy height of the forest at these two sites were similar; midstory trees (< 12.7 cm DBH) in this area averaged 32 m in height and overstory trees (≥ 12.7 cm diameter at DBH) averaged 53 m in height. The two sites were at similar elevations (283 m and 279 m, respectively). We sampled the macroarthropod community concurrently at each site over multiple nights in 2010 using 10 W blacklight traps (Universal Light Trap, Bioquip Products, Gardena, CA), focusing on Lepidoptera. A single blacklight trap was placed on the ground at each site and operated from dusk to dawn. Surveys were carried out from 6 May – 15 August and the two sites were each visited five times (n = 10 trap-nights). As per recommendations by Yela and Holyoak (1997), survey nights were fair with temperatures ≥ 16°C at sunset, no precipitation, and low wind. A cotton tuft soaked in ethyl acetate was placed 2013 THE GREAT LAKES ENTOMOLOGIST 3 in each trap to subdue trapped insects. Following a trap night, specimens were sorted and placed in cold storage (4°C) for identification in the laboratory. Specimens were enumerated and Lepidoptera identified to species (or genus in rarer cases) using Covell (2005), Holland (1903), and reference collections at the University of Kentucky. Voucher specimens were incorporated into existing reference collections. Reports of food habits from Covell (2005) were noted for the most common species as well as those unique to either collection site. Our classification of noctuiods follows that of LaFontaine and Schmidt (2010). We focused our efforts on macrolepidoptera and those microlepidoptera with wings ≥ 20 mm (i.e., some Oecophoridae, Yponomeutidae, Tortricidae, Megalopygidae, Limacodidae, and Pyralidae); thus, our study is not an exhaustive assessment of the assemblage. We calculated Jaccard’s and Sørensen’s coefficients of similarity to report the degree of species overlap between the two sites (Southwood 1978). While limited replication across study sites prohibited a statistical test for differences in Lepidopteran occurrence between the deciduous and hemlock sites, repeated sampling permitted testing for seasonal differences in our data. We tested for differences in the abundance and species richness of Lepidoptera captured per trap using the Wilcoxon rank-sum test in SAS (v. 9.1). Additionally, we used EstimateS (v. 8.2) to generate ICE (Lee and Chao 1994) and Chao 2 (Chao 1987) species richness estimations for comparisons between survey sites. Estimations were based on 1,000 randomizations (Summerville and Crist 2005).

Results A total of 1,020 Lepidoptera was captured. Of these, 559 individuals were captured from the hemlock site and 461 individuals were captured from the deciduous site. Fewer Lepidoptera were captured early in May and June compared to later July and August (Fig. 1).

Figure 1. Mean abundance of Lepidoptera ± SE collected with blacklight traps from a hemlock-dominated and a deciduous-dominated forest in the Daniel Boone National Forest, Kentucky, during May-August of 2010. Fewer Lepidoptera were captured in May and June versus July and August (W = 16.0, df = 9, P = 0.02). 4 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

We identified 733 Lepidoptera (72%) beyond the family level, representing ≥ 137 species and 18 families (Table 1). Of these, ≥ 107 species were identified from the hemlock site and ≥ 85 were identified from the deciduous site. No difference in species richness was detected between the early (May and June) and late (July and August) portions of the season (W = 21.0, df = 9, P > 0.05). The ICE estimates of richness (mean ± std. dev.) were 245 ± 15 and 189 ± 12 species for the hemlock and deciduous forest, respectively. Chao 2 estimates of richness were more conservative, with 216 ± 32 and 162 ± 25 species for the hemlock and deciduous sites, respectively. Geometridae and Erebidae were the most speciose families overall, followed distantly by Noctuidae, Notodon- tidae, and Limacodidae (Fig. 2). These five families comprised > 75% of the Lepidoptera we identified. There were 58 species from 13 families ubiquitous to both sites. Most were in the Geometridae and Erebidae (14 and 13 species, respectively). Sørensen’s coefficient revealed 60% overlap in species similarity, whereas Jaccard’s coefficient revealed a 43% overlap between sites. Clemensia albata Packard (Erebidae) was the most abundant species trapped (n = 40); larvae are fairly ubiquitous in mesic woods and feed on tree lichens. Thirty-four individuals were captured at the hemlock site, whereas only 6 were captured at the deciduous site. The second most commonly encountered species was Palpita magniferalis (Walker) (Pyralidae) (n = 38). Though larvae feed on ash (Fraxinus spp.), adults were captured in approximately equal numbers from both sites. There were 49 species across 10 families that were unique to the hemlock site; most were in the Erebidae, Geometridae, and Noctuidae (14, 12, and 12 species, respectively). A total of three geometrid species known to specialize on hemlocks and other conifers were unique in their collection at the hemlock site; these included, Lambdina fiscellaria (Guenée), Macaria fissinotata (Walker), and M. signaria (Hübner). A number of erebid detritivores were likewise unique to the hemlock site: Idia spp., Scoleocampa liburna (Geyer), Tetanolita mynesalis (Walker), Zanclognatha cruralis (Guenée). Twenty seven species across 11 families were unique to the deciduous site; most were in the Erebidae and Geometridae (14 and 12 species, respectively). Neither unique detritivores, nor conifer specialists were noted in collections at the deciduous site. Family and species composition shifted over the course of our surveys. Our trapping effort in May – June yielded 81 species, primarily Geometridae and Erebidae (27 and 15 species, respectively). There were three commonly captured species/genera: the geometrid genus, Iridopsis (n = 23), as well as the pyralid species, Desmia funeralis (Hübner) (n = 13) and P. magniferalis, (n = 10). Iridopsis spp. feed on a wide variety of herbaceous and woody plants. D. funeralis feeds on the eastern redbud (Cercis canadensis L., Fabaceae) and native grape (Vitis spp.). During July – August, 104 species were captured. While most of these were again Erebidae or Geometridae (27 and 23 species, respectively), the Notodontidae and Noctuidae were also common (14 and 13 species, respectively). This late season trapping yielded many common species/ genera (n ≥ 10 individuals) from eight families, including: Drepanidae: Drepana arcuata Walker, P. magniferalis; Erebidae: Halysidota tessellaris (J. E. Smith); Geometridae: Biston betularia (L.), Caripeta divisata Walker, C. albata, Iridop- sis sp., Macaria sp; Limacodidae: Lithacodes sp., Nadata gibbosa (J. E. Smith); Noctuidae: Marimatha nigrofimbria(Guenée), Polygrammate hebraeicum Hüb- ner; Notodontidae: Heterocampa obliqua Packard, Peridea basitriens (Walker); Pyralidae: D. funeralis; Saturnidae: Anisota stigma (F.), Datana angusii Grote and Robinson, Dryocampa rubicunda (F.). Nearly twice as many species were exclusively captured during July – August (n = 63) versus those exclusively captured during May – June (n = 34). 2013 THE GREAT LAKES ENTOMOLOGIST 5

Table 1. A checklist of Lepidoptera collected in blacklight traps from a hemlock-dominated and a deciduous-dominated forest in the Daniel Boone National Forest, Ken- tucky, during May-August of 2010. Values are the sum of individuals captured within specified months. Asterisks denote capture in both forest types, whereas a superscript denotes capture in only hemlock (h) or deciduous forest (d).

Taxon Hemlock Deciduous

May-June July-Aug. May-June July-Aug.

APATELODIDAE Apatelodes torrefacta (J. E. Smith) * 1 1 3 6 Olceclostera angelica (Grote) * 1 1

DREPANIDAE Drepana arcuata Walker * 11 3 7 Eudeilinia herminiata (Guenée) h 2 Oreta rosea (Walker) * 2 1 1

ELACHISTIDAE Antaeotricha spp. h 1 Antaeotricha schlaegari (Zeller) * 7 5

EREBIDAE Allotria elonympha (Hübner) * 2 3 Apantesis sp. * 1 1 2 Cisseps fulvicollis (Hübner) d 5 Cisthene sp. h 3 Clemensia albata Packard * 2 32 5 1 Crambidia pallida Packard d 2 Crambidia sp. h 1 Cycnia tenera Hübner * 2 2 Dasychira sp. * 1 8 Euparthenos nubilis (Hübner) h 1 Halysidota tessellaris (J. E. Smith) * 3 11 4 Haploa clymene (Brown) * 3 6 Hypena scabra (F.) h 1 1 Hypoprepia fucosa Hübner h 2 Idia sp. h 1 Orgyia leucostigma (J. E. Smith) * 2 2 1 Palthis sp. h 1 Pangrapta decoralis Hübner h 3 Panopoda carneicosta Guenée h 1 Parallelia bistriaris Hübner h 1 Ptichodis herbarum (Guenée) d 1 Pygarctia spraguei (Grote) h 4 Pyrrharctia isabella (J. E. Smith) * 1 2 2 Scoleocampa liburna (Geyer) h 2 5 Spilosoma congrua Walker * 3 1 1 Spilosoma sp. h 3 Spilosoma virginica (F.) * 2 2 Tetanolita mynesalis (Walker) h 1 Virbia opella (Grote) * 1 1 2 Virbia sp. * 5 2 Zale sp. h 1 Zanclognatha cruralis (Guenée) h 3 6 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Continued.

Taxon Hemlock Deciduous

May-June July-Aug. May-June July-Aug.

GEOMETRIDAE Biston betularia (L.) * 17 1 Caripeta divisata Walker * 1 12 Costaconvexa centrostrigaria (Wollaston) h 4 Dyspteris abortivaria (Herrich-Schäffer) * 2 1 5 Ecliptoptera atricolorata (Grote & Robinson) * 2 1 Epimecis hortaria (F.) * 2 3 1 Eubaphe mendica (Walker) d 1 Eulithis diversilineata (Hübner) * 2 1 6 Eutrapela clemataria (J. E. Smith) * 3 4 2 Euchlaena amoenaria (Guenée) * 1 1 Euchlaena pectinaria (Dennis & Schiffermüller) h 3 Glena cognataria (Hübner ) h 1 Glena cribrataria (Guenée) h 3 1 Iridopsis sp. * 8 12 15 Lambdina fervidaria (Hubner ) * 4 5 4 Lambdina fiscellaria(Guenée) h 2 Macaria sp. h 15 Macaria fissinotata(Walker) h 4 1 Macaria ocellinata (Guenée) d 1 Macaria promiscuata Ferguson * 1 1 5 Macaria signaria (Hübner) h 2 Metarranthis angularia Barnes & McDunnough d 1 Metarrhanthis hypochraria (Herrich-Schäffer) d 1 Metarrhanthis indeclinata (Walker) h 1 Nemoria bistriaria Hübner * 6 2 Plagodis alcoolaria (Guenée) d 1 Plagodis fervidaria (Herrich-Schäffer) d 3 Plagodis phlogosaria (Guenée) d 1 1 Plagodis serinaria Herrich-Schäffer h 3 Prochoerodes lineola (Drury) h 1 3 Probole amicaria (Herrich-Schäffer) * 1 4 Speranza coortaria (Hulst) * 3 1 Tetracis sp. d 2 Tetracis cachexiata Guenée h 4 Tetracis crocallata Guenée h 2 Trichodezia albovittata (Guenée) d 1

LASIOCAMPIDAE Artace cribraria (Ljungh) d 3

LIMACODIDAE Apoda biguttata (Packard) * 1 3 Apoda y-inversum (Packard) * 2 1 Euclea delphinii (Boisduval) * 1 1 2 2013 THE GREAT LAKES ENTOMOLOGIST 7

Table 1. Continued.

Taxon Hemlock Deciduous

May-June July-Aug. May-June July-Aug.

Lithacodes sp. h 18 Lithacodes fasciola (Herrich-Schäffer) * 4 4 Natada nasoni (Grote) * 1 2 Parasa chloris (Herrich-Schäffer) d 2 Prolimacodes badia (Hübner) h 8 Torticidia flexuosa(Grote) d 2

MEGALOPYGIDAE Lagoa crispata (Packard) h 3 8

NOCTUIDAE Abagrotis alternata (Grote) h 1 Acronicta americana (Harris) h 1 Acronicta haesitata (Grote)d 6 Acronicta inclara J. B. Smith h 1 Acronicta funeralis Grote & Robinson h 2 Agrotis ipsilon (Hufnagel) h 1 Cosmia calami (Harvey) h 1 Eudryas grata (F.) * 1 1 1 Feltia jaculifera (Guenée) h 1 Lacinipolia renigera (Stephens) d 1 Leuconycta diptheroides (Guenée) h 1 Marimatha nigrofimbria (Guenée) * 9 1 1 Morrisonia confusa (Hübner) d 1 Orthodes crenulata (Butler) h 1 Polygrammate hebraeicum Hübner h 11 Pseudeustrotia carneola (Guenée) h 1 Spodoptera dolichos (F.) h 1 Xestia dolosa Franclemont * 5 2

NOLIDAE Baileya australis (Grote) * 3 1 1

NOTODONTIDAE Cerura scitiscripta Walker h 2 Datana angusii Grote & Robinson * 3 5 5 7 Datana contracta Walker * 3 2 1 2 Datana ministra (Drury) d 3 Datana perspicua Grote & Robinson h 3 Heterocampa obliqua Packard * 3 4 2 14 Lochmaeus bilineata (Packard) h 1 Lochmaeus manteo Doubleday h 1 Macrurocampa marthesia (Cramer) d 2 4 Nadata gibbosa (J. E. Smith) * 3 6 4 8 Nerice bidentata Walker h 1 Oligocentria lignicolor (Walker) d 3 Peridea basitriens (Walker) * 2 4 3 14 Symmerista albifrons (J. E. Smith) * 4 2 2

8 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Continued.

Taxon Hemlock Deciduous

May-June July-Aug. May-June July-Aug.

OECOPHORIDAE Psilocorsis spp. * 5 2 3

PYRALIDAE Blepharomastix ranalis (Guenée) d 1 Desmia funeralis (Hübner) * 1 25 12 Galleria mellonella (L.) h 3 Palpita magniferalis (Walker) * 3 14 7 14 Pantographa limata (Grote & Robinson) * 1 6 2 1

SATURNIIDAE Actias luna (L.) d 2 Anisota stigma (F.) * 3 10 Automeris io (F.)* 2 1 Callosamia angulifera (Walker) d 1 Dryocampa rubicunda (F.) * 1 4 12 Eacles imperialis (Drury) * 1 2

SPHINGIDAE Ceratomia undulosa (Walker) * 1 1 Deidamia inscripta (Harris) d 1 Paonias excaecatus (J. E. Smith) * 2 2

THYATIRIDAE Pseudothyatira cymatophoroides (Guenée) d 1

TORTRCIDAE Argyrotaenia alisellana (Robinson) h 1 Clepsis melaleucana (Walker) d 1

YPONOMEUTIDAE Atteva aurea (Fitch) * 2 1 2013 THE GREAT LAKES ENTOMOLOGIST 9

Figure 2. Percent composition of lepidopteran families collected with blacklight traps from a hemlock-dominated and a deciduous-dominated forest in the Daniel Boone National Forest, Kentucky, during May-August 2010. 10 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Discussion Our data demonstrate that the presence of eastern hemlock contributes significantly to the overall lepidopteran diversity of Appalachian forests. While our species richness estimates indicate that our survey was far from comprehensive, it is clear that both hemlock and deciduous forests possess rich lepidopteran assemblages. It is notable that nearly twice as many species were unique to our hemlock site than our deciduous site and that a number of these species were either detritivores or conifer specialists. This implies that on a limited spatial scale hemlock-dominated habitats may possess a suite of Lepidoptera not found in habitats with a deciduous overstory. This supports results from other studies. In its role as a foundation species in eastern North America, eastern hemlock maintains a distinct community of benthic, riparian and canopy arthropod associates. Over 200 insect and 33 mite species are reportedly associated with hemlock in central Appalachia (Wallace and Hain 2000, Buck et al. 2005, Dilling et al. 2007, Turcotte 2008, J. K. Adkins and L. K. Rieske unpublished data), and hemlocks support greater spider abundance, richness, diversity and evenness than do deciduous canopies (Mallis and Rieske 2011), including several species unique to hemlock (Aiken and Coyle 2000). Our results must be interpreted with caution, however, as many Lepidoptera are strong fliers and their appearance in traps associated with a given forest type may not indicate resident populations. Dodd et al. (2012) found lepidopteran abundance and diversity in the hardwoods of Central Appalachia to be higher after May. While we found that Lepidoptera were more abundant later in the sampling period, we failed to detect differences in overall species richness. Even so, our data demonstrate a high amount of turnover in the occurrence of species over the sampling period. Further, we observed more common species in July and August, suggesting a more even lepidopteran assemblage later in the season. Our data underpin the recommendations by Summerville and Crist (2005), that in order to capture much of the local variation in lepidopteran occurrence studies must span a wide window of time in order to adequately inventory species. Our study forms a foundation for understanding species richness patterns of Lepidoptera in hemlock forests in North America and is a useful baseline for comparisons of richness and diversity post invasion by the hemlock woolly aldegid. The adelgid is expanding its geographic range at an alarming rate, and hemlock forests in eastern North America are at risk. Given the potential for direct and indirect impacts of this invasion on Lepidoptera and other arthropods (Dilling et al. 2009), baseline knowledge of species occurrence is essential. Such knowledge can help land managers assess impacts and mitigate effects of current and future invasions on forest health and forest sustainability.

Acknowledgments The authors thank T. Culbertson for technical assistance. This project was supported in part by McIntire Stennis funds provided through the University of Kentucky’s College of Agriculture, and is published as Kentucky Agricultural Experiment Station publication number 12-08-054.

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Buck, S. L., P. Lambdin, D. Paulsen, J. Grant, and A. Saxton. 2005. Checklist of insect species associated with eastern hemlock in the Great Smoky Mountains Na- tional Park and environs. Tennessee Academy of Science 80: 1-10. Chao, A. 1987. Estimating the population size for capture–recapture data with unequal catchability. Biometrics 43: 783-791. Covell, C. V. 2005. A field guide to moths of Eastern North America: Special Publication Number 12. Virginia Museum of Natural History, Martinsville, Virginia. Dilling, C., P. Lambdin, J. Grant, and L. Buck. 2007. Insect guild structure associated with eastern hemlock in the Southern Appalachians. Environmental Entomology 36: 1408-1414. Dilling, C., P. Lambdin, J. Grant, and R. Rhea. 2009. Community response of insects associated with eastern hemlock to Imidacloprid and horticultural oil treatments. Environmental Entomology 38: 53-66. Dix, M. E., R. J. Johnson, M. O. Harrell, R. M. Case, R. J. Wright, L. Hodges, J. R. Brandle, M. M. Shoeneberger, R. L. Fitzmaurice, L. J. Young, and K. G. Hubbard. 1995. Influences of trees on abundance of natural enemies of insect pests: a review. Agroforestry Systems 29: 303-311. Dodd, L. E., M. J. Lacki, and L. K. Rieske. 2008. Variation in moth occurrence and implications for foraging habits of Ozark big-eared bats. Forest Ecology and Man- agement 255: 3866-3872. Dodd, L. E., M. J. Lacki, and L. K. Rieske. 2011. Habitat associations of Lepidoptera in the Ozark Mountains of Arkansas. Journal of the Kansas Entomological Society 84: 271-284. Dodd, L. E., M. J. Lacki, E. R. Britzke, D. A. Buehler, P. D. Keyser, J. L. Larkin, A. D. Rodewald, T. B. Wigley, P. B. Wood, and L. K. Rieske. 2012. Forest structure affects trophic linkages: how silvicultural disturbance impacts bats and their insect prey. Forest Ecology and Management 267: 262-270. Godman, R. M., and K. Lancaster. 1990. Tsuga Canadensis, pp. 604-612. In R. M. Burns and B. H. Honkala [eds.], Silvics of North America: Vol.1. USDA Forest Service Agricultural Service. Handbook 654. Washington, D.C. Holland, W. J. 1903. The moth book. Doubleday, Page and Company, New York City, New York. Lacki, M. J., S. K. Amelon, and M. D. Baker. 2007. Foraging ecology of forest bats, pp. 83-127. In Bats in forests: conservation and management (eds. Lacki, M. J., J. P. Hayes, and A. Kurta).John Hopkins University Press, Baltimore, MD. LaFontaine, J. D., and B. C. Schmidt. 2010. Annotated check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico. Zookeys 40: 1–239. Lee, S. M., and A. Chao. 1994. Estimating population size via sample coverage for closed capture–recapture models. Biometrics 50: 88–97. Mallis, R. E., and L. K. Rieske. 2011. Arboreal spiders in eastern hemlock. Environ- mental Entomology 40: 1378-1387. Pohl, G. R., D. W. Langor, J.-F. Landry, and J. R. Spence. 2004. Moths and butterflies (Lepidoptera) of the boreal mixedwood forest near Lac La Biche, Alberta, including new provincial records. The Canadian Field Naturalist 118: 530-549. Rohr, J. R., C. G. Mahan, and K. C. Kim. 2009. Response of arthropod biodiversity to foundation species declines: The case of the eastern hemlock. Forest Ecology and Management 258: 1503-1510. Ross, R., R. Bennett, C. Snyder, D. Smith, J. Young, D. Lemarie. 2003. Influence of eastern hemlock (Tsuga canadensis L.) on fish community structure and function in headwater streams of the Delaware River basin. Ecology of Freshwater Fish 12: 60–65. 12 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Snyder, C., J. Young, D. P. Lemarié, and D. R. Smith. 2002. Influence of eastern hemlock (Tsuga canadensis) forests on aquatic invertebrate assemblages in headwater streams. Canadian Journal of Fish Aquatic Sciences 59: 262–275. Southwood, T. R. E. 1978. Ecological Methods, Second Edition. Chapman and Hall, London. Summerville, K. S., and T. O. Crist. 2001. The species richness of Lepidoptera in a fragmented landscape: a supplement to the checklist of moths in Butler County, Ohio. The Great Lakes Entomologist 34: 93-110. Summerville, K. S., and T. O. Crist. 2005. Temporal patterns of species accumulation in a survey of Lepidoptera in a beech-maple forest. Biodiversity and Conservation 14: 3393-3406. Summerville, K. S. and T. O. Crist. 2008. Structure and conservation of lepidopteran communities in managed forests of northeastern North America: a review. Canadian Entomology 140: 475-494. Summerville, K. S., J. J. Jacquot, and R. F. Stander. 1999. A preliminary checklist of moths in Butler County, Ohio. Ohio Journal of Science 99: 66-76. Thomas, A. W. 2001. Moth Diversity in a Northeastern North American Red Spruce Forest: I. Baseline Study. Information Report, M-X-210E, Canadian Forest Service, Canada. Thomas, A. W., and G. M. Thomas. 1994. Sampling strategies for estimating moth species diversity using a light trap in a northeastern softwood forest. Journal of the Lepidopterists Society 48: 85-105. Tingley, M. W., D. A. Orwig, R. Field, and G. Motzkin. 2002. Avian response to removal of a forest dominant: consequences of hemlock woolly adelgid infestations. Journal of Biogeography 29: 1505-1516. Turcotte, R. M. 2008. Arthropods associated with eastern hemlock. p.61, In B. Onken and R. Reardon (eds.) Fourth Symposium on hemlock woolly adelgid in the eastern United States. USDA Forest Service FHTET-2008-01, Morgantown, West Virginia. Wallace, M. S. and F. P. Hain. 2000. Field surveys and evaluation of native and established predators of the hemlock woolly adelgid (Homoptera: Adelgidae) in the southeastern United States. Environmental Entomology 29: 638-644. Woods, A. J., J. M. Omernik, W. H. Martin, G. J. Pond, W. M. Andrews, S. M. Call, J. A. Comstock, and D. D. Taylor. 2002. Ecoregions of Kentucky (color poster with map, descriptive text, summary tables, and photographs). US Geological Survey, Reston, Virginia (map scale 1:1,000,000). Yela, J. L., and M. Holyoak. 1997. Effects of moonlight and meteorological factors on light and bait trap catches of noctuid moths. Environmental Entomology 2: 1283-1290. Yorks, T. E., J. C. Jenkins, D. J. Leopold, J. R. Dudley, and D. A. Orwig. 2000. Influences of eastern hemlock mortality of nutrient cycling, pp.126-133.In Proceed- ings of the symposium on sustainable management of hemlock ecosystems in eastern North America (eds K. A. McManus, K. S. Shields, and D. R. Souto). General Techni- cal Report NE-267. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. Yorks, T. E., D. J. Leopold and D. J. Raynal. 2003. Effects of Tsuga canadensis mortality on soil water chemistry and understory vegetation: possible consequences of an invasive insect herbivore. Canadian Journal of Forest Research 33: 1525-1537. 2013 THE GREAT LAKES ENTOMOLOGIST 13 Attraction of Agrilus planipennis (Coleoptera: Buprestidae) and Other Buprestids to Sticky Traps of Various Colors and Shapes Toby R. Petrice1, Robert A. Haack1, and Therese M. Poland1

Abstract The family Buprestidae (Coleoptera) contains numerous economically significant species, including the emerald ash borer (EAB), Agrilus planipennis Fairmaire, first discovered in North America in 2002. Effective traps for monitoring spread and population densities of EAB and other buprestids are needed. Studies were conducted in 2008 to test different colors and shapes of sticky traps baited with manuka oil for capturing EAB and other buprestids. Among different trap shapes, an enlarged purple Agrilus-shaped silhouette (15 cm wide × 55 cm all) attached to a white background (40 cm wide × 60 cm tall) captured the most buprestid species compared to purple traps (40 cm wide × 60 cm tall) with or without a single dead EAB adult decoy attached at the trap center. The mean number of buprestid species captured per trap were intermediate on purple traps with 25 dead EAB adult decoys, an enlarged green Agrilus-shaped silhouette (15 cm wide × 55 cm tall) attached to a white background (40 cm wide × 60 cm tall), and an enlarged EAB photograph (15 cm wide × 55 cm tall) on a white background (40 cm wide × 60 cm tall). There were no significant differences detected among the different trap shapes when total number of buprestids captured per trap were compared. However, purple traps with 25 EAB adult decoys captured significantly more EAB per trap compared to enlarged EAB photographs, enlarged purple Agrilus-shaped silhouettes, or purple traps without decoys. In another study, there were no significant differences detected in the mean number of buprestid species, total buprestids, or EAB adults captured per trap among purple, green, or half purple and half green three-sided prism-shaped traps (each side = 40 cm wide × 60 cm tall). Response to different trap shapes and colors varied among some buprestid species and these differences are discussed.

______

The family Buprestidae (Coleoptera) includes numerous economically significant species in North America, especially in the genusAgrilus . Among these is the nonnative invasive species emerald ash borer [EAB, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae)], first discovered in North America in 2002 (Haack et al. 2002). Native to Asia, EAB has become one of the most destructive forest insect pests introduced into North America (Cappaert et al. 2005, Poland and McCullough 2006, Kovacs et al. 2010). Adult EAB oviposit on ash (Fraxinus spp.) trees and likely use both olfactory and visual cues to locate hosts and mates. Research on EAB olfactory cues has focused on host kairomones and possible specific pheromones (Rodriguez- Saona et al. 2006; Crook et al. 2008a, b; Silk et al. 2011; Ryall et al. 2012). Research on visual attraction in EAB has been broad in scope but the number of studies has been limited. Studies have found that EAB adults are attracted

1USDA Forest Service, Northern Research Station, 1407 S. Harrison Rd., Michigan State University, East Lansing, MI 48823. *Corresponding author: (e-mail: [email protected]). 14 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 to the colors purple (Oliver et al. 2004; Francese et al. 2005, 2008) and green (Crook et al. 2009; Francese et al. 2010). In addition, Lelito et al. (2008) found that EAB adults were attracted to dead EAB adults in the field when placed as decoys on ash leaflets. Literature suggests buprestids are very visually oriented. For example, in Australia, Julodimorpha bakewelli (White) (Coleoptera: Buprestidae) males were found to be attracted to glass bottles (370 mL) that resemble the color and texture of J. bakewelli females (Gwynne and Rentz 1983). Domingue et al. (2011) reported that three European Agrilus species were attracted to dead Agrilus adults attached to foliage. Lelito et al. (2011) found similar attraction for Agrilus subcinctus Gory (native) and Agrilus cyanescens Ratzeburg (nonnative) in the United States. In a field study conducted in Bulgaria, Sakalian et al. (1993) found most beetles, including buprestids, to be attracted to yellow or white conical traps, compared to black, blue, green, orange, or red. In contrast, Oliver et al. (2003) conducted a study in the United States and found buprestids to be most attracted to red-colored traps compared to blue, green or white. In the study by Sakalian et al. (1993), 99% of the buprestids captured were in the genera Anthaxia and Acmaeodera, which are species that commonly visit flowers. Oliver et al. (2003) captured primarily Chrysobothris species (78 %) which are usually not associated with flowers. Traps for monitoring buprestid populations are limited primarily to single- colored sticky traps. Currently, detection traps for EAB in the U.S. and Canada consist of large, three-sided (each rectangular side = 36 cm wide × 60 cm tall) purple or green corrugated plastic prism-shaped traps (Francese et al. 2010). The outer surface of these traps is coated with insect glue to entrap landing EAB adults. McCullough et al. (2011) and Poland et al. (2011) found that a multicomponent trap was more effective for capturing EAB at low beetle densities compared to three-sided prism traps. More effective traps for monitoring spread and population densities of EAB and other buprestids are needed. In 2008, we conducted studies to evaluate the attraction of buprestids, including EAB, to sticky traps of different shapes or colors, as well as traps with dead EAB adults added as decoys, or with a photograph of an enlarged EAB adult or its silhouette.

Materials and Methods Studies took place at four sites in Ingham County, Michigan: Legg Park (Lat 42.69 N, Long -84.38 W), Ferguson Park (Lat 42.71 N, Long -84.43 W), Wonch Park (Lat 42.71 N, Long -84.43 W), and a woodlot in Dansville, MI (Lat 42.55 N, Long -84.32 W). The three parks consisted of open grassy areas surrounded by mature trees of mixed hardwood species including green ash, Fraxinus pennsylvanica Marsh. The woodlot in Dansville was composed of mixed hardwoods including green and white ash (Fraxinus americana L.). At each site, EAB populations were high, as evidenced by abundant dead ash trees with EAB exit holes and live ash trees with numerous woodpecker feeding sites in the bark that were created while foraging for EAB larvae. Trap shape study. In the first visual trapping experiment there were six treatments (Fig. 1). Three of the treatments consisted of two-sided-rectangular pieces of dark purple corrugated plastic [stock color manufactured by Coroplast, Inc., Vanceburg, KY; see Francese et al. (2010) for reflectance values], 40 cm wide × 60 cm tall, with either A) 25 dead EAB adults (sex not determined) per side attached with a cyanoacrylate adhesive (Loctite® Super Glue, Henkel Corporation, Westlake, OH) in 5 rows of 5 adults each spaced over the entire surface of each side of the trap to serve as decoys, B) 1 dead EAB adult (sex not determined) attached in the center of each side of the trap, or C) no EAB adult decoys (control). The fourth treatment was a 15 cm wide × 55 cm tall enlarged photograph of an EAB adult in dorsal view printed on a white background (modified from original photograph taken by Klaus Bolte, Canadian Forest Service, 2013 THE GREAT LAKES ENTOMOLOGIST 15 A B C

D E F

Figure 1. Traps used for trap shape study including: (A) purple rectangle with 25 dead emerald ash borer (EAB) adults, (B) purple rectangle with 1 dead EAB adult attached to center, (C) purple rectangle with no EAB adults (control), (D) enlarged EAB photograph on white background (modified from original photograph taken by Klaus Bolte, Canadian For- est Service, Natural Resources Canada, now retired), (E) purple Agrilus silhouette on white background, and (F) green Agrilus silhouette on white background. Traps were 40 cm wide ´ 60 cm tall flat panels with the same treatment duplicated on each side.

Natural Resources Canada, now retired). The last two treatments consisted of a single 15-cm-wide × 55-cm-tall dark purple (Coroplast stock color)) or light green [”young ash green” manufactured by Coroplast, Inc., Vanceburg, KY; see Francese et al. (2009) for reflectance values] silhouette of an adultAgrilus beetle viewed dorsally (traced from an enlarged EAB photograph) that was cut from corrugated plastic. Agrilus silhouettes and EAB photographs were glued to the center of both sides of a 40 cm wide × 60 cm tall white foam poster board with construction adhesive (Liquid Nails® Heavy Duty Construction Adhesive, Liquid Nails Adhesive, Strongsville, OH). The rationale for the latter three traps types was that EAB and other buprestids are visually attracted to conspecifics (Lelito et al. 2008, Domingue et al. 2011) and enlarged silhouettes might be visible to buprestids from greater distances. Three replicates of each treatment were placed at Legg Park, 3 at Ferguson Park, and 4 at Dansville for a total of 10 replicates. Traps were deployed 13 June 2008 and removed on 12 August 2008. Trap color study. In the second visual trapping experiment, purple and green three-sided-prism traps were compared (Fig. 2). Traps consisted of a 108 cm ´ 60 cm rectangular piece of dark purple (Coroplast stock color) or light green (”young ash green”) corrugated plastic folded twice perpendicular to the long edge to make three 36 cm ´ 60 cm sides. The free ends were fastened together using plastic ties. Treatments included traps that were all purple, 16 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 A B

C D

Figure 2. Traps used for trap color study including: (A) all purple prism, (B) all green prism, (C) purple top and green bottom prism, and (D) green top and purple bottom prism. Traps were three-sided with the same treatment on each side. Traps were 36 cm-wide ´ 60 cm-tall on each side. all green, purple on the top half and green on the bottom half, or green on the top half and purple on the bottom half. For traps with both purple and green, construction adhesive was used to attach green plastic over the top- or bottom half of the purple plastic. Five replicates each were placed at Legg Park and Wonch Park for a total of 10 replicates. Traps were deployed 13 June 2008 and removed on 12 August 2008. Trap setup, monitoring, and data analyses. For both studies, all traps were suspended from rebar poles (1.25 cm diam.) approximately 2 m above the ground. The bottom of the rebar pole was inserted into the ground and the top had a 90º bend approximately 0.3 m long, where each trap was attached. Traps were spaced 15 m apart and situated on a forest edge a minimum of 15 m from any ash tree. 2013 THE GREAT LAKES ENTOMOLOGIST 17 All surfaces of each trap were painted with clear Pestick™ insect glue (Hummert International, Earth City, MO). Pestick™ was carefully applied to those traps with the dead EAB adults to avoid covering the individual EAB decoys. Since EAB was the primary species targeted for both trapping experiments, all traps were baited with manuka oil (manufactured by Coast Biologicals, Ltd. Auckland, NZ, distributed by Merchant Ag/Response-Ohio, Inc., Chagrin Falls, OH), a steam distillate from the New Zealand tea tree, Leptospermum scoparium J.R. and G. Forst. (Myrtaceae), that has been found to enhance attraction of EAB to sticky traps (Crook et al. 2008b). The manuka oil was dispensed from a cluster of five 0.4-ml polyethylene snap-cap tubes (Fisher Scientific, Pittsburgh, PA) at a total combined release rate of 10 mg/d. Buprestids were collected from traps every 1–3 d from 14 June through 15 July 2008 (to include EAB peak flight), then collected every 1–2 wk through 12 August 2008. Samples were frozen until they could be cleaned and beetles identified in the laboratory. Hexane was used to remove Pestick™ from beetles prior to identification. Buprestids were identified to species using keys in Fisher (1928), Wellso et al. (1976), Wellso and Manley (2007), and MacRae (2003). Sex of each beetle was determined using morphological characters or dissection of genitalia when required for some species. Female members of the Agrilus otiosus species-group and a few related Agrilus species cannot be distinguished from one another morphologically (Fisher 1928, MacRae 2003). For this paper, therefore, these females were pooled and referred to as female A. otiosus-relatives. Male Agrilus otiosus-relatives captured in our study included: Agrilus arcuatus (Say), Agrilus defectus Leconte, Agrilus masculinus Horn, Agrilus otiosus Say, and Agrilus transimpressus Fall. We pooled all male A. otiosus-relatives for statistical analyses which enabled comparison with females, and because males of most species were represented by only a few specimens. In addition, female Agrilus celti Knull and Agrilus egenus Gory cannot be distinguished from each other morphologically (Fisher 1928), and are referred to as A. celti + A. egenus in this paper. For statistical analyses, we chose to pool males of both of these species as well. The percentage of beetles captured per trap was calculated as the number captured by an individual trap expressed as a percentage of the total number captured by all traps within each replicate for the entire flight season. Per- centages were calculated for different buprestid groups, individual species, or sexes relative to the total number of the respective buprestid group, species, or sex within each replicate. Replicates that did not capture any specimens of a given buprestid group, species, or sex were excluded from that particular analysis. Responses to different traps were analyzed by genus, species, and sex when adequate numbers existed, i.e., >25 total representatives of a buprestid group or species. Percentage data were normalized using arcsine square root transformations before analyses. The number of species captured was normalized using Log10 transformations before analyses. Data were analyzed with PROC GLM (SAS 2008). Means separation was performed with Least Squared Differences when analyses were significant at theP < 0.05 level.

Results Trap shape study. Overall, 776 buprestids were captured on the different shaped traps, representing 4 genera and at least 28 species (Table 1). Of these, 672 individuals were species of Agrilus, representing at least 17 species of which 530 individuals were EAB. The next two most abundant buprestid genera captured were Chrysobothris (89 individuals and at least 7 species) and Anthaxia (12 individuals and at least 3 species). The mean percentage of all buprestids combined per trap did not vary significantly among trap shapes (Fig. 3a; significance values are given in Figs. 3-6). However, there was a significant difference in the mean number of buprestid species captured per trap among the 18 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 1 5 1 1 1 (6) 1(0) 0(1) 0(3) 2(0) 1(4) 0(3) 0(1) 2(2) 0(7) 1(1) (33) 0(1) 1(5) 4(9) 0(3) 22(32) 197(333) captured Total buprestids 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (0) (1) 1(0) 0(1) 0(1) 1(0) 0(3) 41(60) Purple control 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (7) (0) 0(2) 0(1) 1(0) 0(3) 3(7) 1(2) 46(83) 25 EAB Purple with 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 (0) (1) (0) 0(1) 1(2) 0(1) 0(4) 36(59) 1 EAB Purple with 0 0 0 1 0 0 0 1 0 1 0 0 0 0 0 0 0 (3) (1) 1(3) 2(1) 7(25) 13(12) Enlarged Trap type EAB photo

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 (4) (1) 0(1) 0(2) 5(4) 34(52) Purple silhouette

0 0 2 0 4 0 0 0 0 0 1 (3) 0(1) 0(2) 0(1) 0(1) 0(3) 0(1) (18) 0(3) 0(2) 3(7) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) 33(54) Green silhouette

2 2

1 silhouettes (15 cm wide × 55 tall) on white backgrounds; enlarged EAB photographs cm) attached to

1 Barter&Brown

1 Gory Fall males Leconte females Agrilus (Say) Gory Gory Fairmaire liragus Horn males (Randall)

Ratzeburg Agrilus Anthaxia (Weber) Lacordaire Leconte males Saunders Say males -relative females (Say) Gory-males Knull males Table1. Number of male and female buprestids captured in Ingham County, Michigan, June-August 2008, on sticky traps with enlarged purple or green 40 cm wide × 60 tall white background; or 0, 1, 25 dead EAB adults attached to purple rectangles (see Fig.1). Traps (10 replicates per treatment) were two-sided, coated with Pestick™ insect trapping glue, and baited manuka oil mg/d). Species Agrilus anxius A. bilineatus A. celti A. celti + egenus A. cyanescens A. defectus A. egenus A. granulatus A. lecontei A. masculinus A. obsoletoguttatus A. otiosus A. otiosus A. planipennis A. politus A. sulcicollis A. transimpressus A. vittaticollis Unidentified Anthaxia expansa An. viridicornis An. viridifrons Unidentified 2013 THE GREAT LAKES ENTOMOLOGIST 19 0(1) 0(1) 1(3) 0(3) 0(3) 2(2) 0(10) 13(25) 11(17) 267(509) captured Total buprestids 0 0 0 0 0 0(1) 1(2) 0(3) 1(1) 45(73) Purple control 0 0 0 0 0 0(1) 1(0) 0(1) 0(2) 52(109) 25 EAB Purple with 0 0 0 0 0 2(4) 0(2) 2(1) 1(0) 43(75) 1 EAB Purple with 0 0 0 5(8) 0(1) 0(2) 0(1) 3(8) 1(2) 35(67) Enlarged Trap type EAB photo

0 0 0 0(1) 0(2) 0(2) 0(4) 5(4) 3(10) 47(87) Purple silhouette

0 0 0 0 0 0 2(0) 0(1) 0(1) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀) 45(98) Green silhouette Harold Gory&Laporte species group or a close relative. Females in this group cannot be distinguished from one another. See MacRae (2003) for a species group or a close relative. Females in this (Fabricius) Chrysobothris Say Melsheimer Melsheimer (Olivier) Wellso&Manley A. otiosus Females of these two species cannot be distinguished from one another. In the Table1. Continued. Species Chrysobothris adelpha C. femorata C. quadriimpressa C. rugosiceps C. sexsignata C. shawnee C. viridiceps Unidentified Dicerca lurida Total buprestids captured 1 2 complete list of species in this group. Figure 6. Mean number of Buprestidae or Agrilus species captured per trap in Ingham County, Michigan, during June-August 2008 on three-sided prism sticky

20 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 3. Mean percentage of Buprestidae (a), and the most common buprestid genera and species (b-f),captured per trap in Ingham County, Michigan, during June-August 2008 on sticky traps with enlarged purple or green Agrilus silhouettes (15 cm wide × 55 cm tall) on white backgrounds; enlarged EAB photographs on white backgrounds (15 cm × 55 cm); or 0, 1, or 25 dead EAB adults attached to a purple background. Traps (40 cm wide × 60 cm tall) were two-sided and baited with manuka oil (5 mg/d). Means within each buprestid group and sex followed by a different letter are significantly different at theP < 0.05 level (LSD, N = 10). Figure 6. Mean number of Buprestidae or Agrilus species captured per trap in Ingham County, Michigan, during June-August 2008 on three-sided prism sticky

2013 THE GREAT LAKES ENTOMOLOGIST 21

Figure 4. Mean number of Buprestidae (a), Agrilus (b), or Chrysobothris (c), species captured per trap in Ingham County, Michigan, during June-August 2008 on traps with enlarged purple or green Agrilus silhouettes (15 cm wide × 55 cm tall) on white backgrounds; enlarged EAB photographs (15 cm × 55 cm) on white backgrounds; or 0, 1, or 25 dead EAB adults attached to a purple background. Traps (40 cm wide × 60 cm tall) were two-sided and baited with manuka oil (10 mg/d). Means within each buprestid group followed by a different letter are significantly different at theP < 0.05 level (LSD, N = 10). 22 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 5. Mean percentage of Buprestidae (a), and the most common genera and species (b-f), captured per trap in Ingham County, Michigan, during June-August 2008 on three- sided prism sticky traps (each side = 36 cm wide × 60 cm tall) that were purple, green, purple on top and green on the bottom, or green on top and purple on the bottom. All traps were baited with manuka oil (10 mg/d). Means within each buprestid group and sex followed by a different letter are significantly different at theP < 0.05 level (LSD, N = 10). 2013 THE GREAT LAKES ENTOMOLOGIST 23

Figure 6. Mean number of Buprestidae (a), or Agrilus (b), species captured per trap in Ingham County, Michigan, during June-August 2008 on three-sided prism sticky traps (each side = 36 cm wide × 60 cm tall) that were purple, green, purple on top and green on the bottom, or green on top and purple on the bottom. All traps were baited with manuka oil (10 mg/d). No significant differences were detected among treatments. 24 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 different trap shapes (Fig. 4a). The purple Agrilus silhouette traps captured significantly more buprestid species compared to the purple traps with a single EAB decoy and the purple traps without EAB decoys, while the mean number of buprestid species captured by the remaining trap types was intermediate between Agrilus silhouette and purple traps (Fig 4a). The mean percentage of all female Agrilus captured per trap, and for both sexes combined, was significantly higher for the purple traps with 25 EAB decoys and the green Agrilus silhouette traps compared to purple traps without EAB decoys and the enlarged EAB photograph traps (Fig. 3b). The mean percentage of combined Agrilus or female Agrilus captured on other trap types was intermediate. The mean percentage of all male Agrilus combined captured per trap did not vary significantly among trap types, but followed a similar trend as female and combined Agrilus in absolute terms. The mean number of Agrilus species captured per trap was significantly higher on the green Agrilus silhouette traps compared to the purple traps with one EAB decoy, purple traps without EAB decoys, and the enlarged EAB photograph traps (Fig. 4b). Purple traps with 25 EAB decoys or the purple Agrilus silhouette traps captured an intermediate number of Agrilus species. The mean percentage of Chrysobothris species combined per trap also varied significantly among treatments (Fig. 3c). Overall, the purple Agrilus silhouette traps captured the highest mean percentage of Chrysobothris males, females and both sexes combined, green Agrilus silhouette traps tended to capture the lowest mean percentages, and the remaining trap types were intermediate (Fig. 3c). Purple Agrilus silhouette traps captured significantly more species of Chrysobothris than did green Agrilus silhouette traps, purple traps with 25 EAB decoys, or purple rectangle traps with no EAB decoys. The remaining treatments captured an intermediate number of Chrysobothris species (Fig. 4c). The mean percentage of EAB males captured per trap was significantly lower on the enlarged EAB photograph traps compared to purple traps with 25 EAB decoys, with 1 EAB decoy, and purple or green Agrilus silhouette traps (Fig. 3d). Purple traps without EAB decoys captured an intermediate percentage of EAB males. The mean percentage of EAB females captured per trap was highest on purple traps with 25 EAB decoys and lowest on the enlarged EAB photograph traps (Fig. 3d). Purple traps with one EAB decoy, green or purple Agrilus silhouette traps, and purple traps without an EAB decoy captured an intermediate percentage of EAB females. The mean percentage of total EAB of both sexes combined followed a similar trend as for EAB females; females dominated the trap catch. The mean percentage of Agrilus anxius Gory males captured per trap, as well as the percentage of both sexes combined, was significantly higher on the enlarged EAB photograph traps compared to purple traps with 25 EAB decoys, 1 EAB decoy, or no EAB decoys (Fig. 3e). The mean percentage of A. anxius males and both sexes combined per trap on purple or green Agrilus silhouette traps was intermediate. The mean percentage of A. anxius females captured per trap did not vary significantly among trap types but followed a similar trend to A. anxius males in absolute terms. Four species of male A. otiosus-relatives were captured on the different trap types, of which A. masculinus represented almost 56% (Table 1). The mean percentage of A. otiosus-relatives that were males, females, or for both sexes combined was significantly higher on greenAgrilus silhouette traps compared to the other traps tested (Fig. 3f). Trap color study. Overall, 1,087 buprestids were captured in the trap color study, representing four genera and at least 20 species (Table 2). Of the total number of buprestids captured, 1,076 were Agrilus species that represented 15 species of which 296 were EAB (Table 2). The mean percentage of buprestids 2013 THE GREAT LAKES ENTOMOLOGIST 25

Table 2. Number of male and female buprestids captured in Ingham County, Michi- gan, June-August 2008, on three-sided prism sticky traps (each side = 36 cm wide × 60 cm tall) that were purple, green, purple on top and green on bottom, or green on top and purple on the bottom (see Fig. 2). Traps (10 replicates per treatment) were coated with Pestick™ insect trapping glue and baited with manuka oil (10 mg/d).

Trap color

Green Purple All over over All Total green purple green purple buprestids Species ♂(♀) ♂(♀) ♂(♀) ♂(♀) ♂(♀)

Agrilus anxius Gory 2(1) 0(1) 0(1) 0(2) 2(5) A. arcuatus (Say) 2 1 0 0 0 1(0) A. bilineatus (Weber) 0 0 0(1) 0 0(1) A. celti Knull males1 0 4 3 0 7 A. celti + egenus females1 (13) (4) (9) (0) (26) A. cyanescens Ratzeburg 1(5) 0(2) 0(3) 0 1(10) A. egenus Gory males1 4 1 2 0 7 A. fallax Say 0 0 0(1) 0 0(1) A. lecontei Saunders 2(12) 1(2) 3(6) 0(2) 6(22) A. masculinus Horn males2 65 24 56 1 146 A. obsoletoguttatus Gory 0(2) 0(1) 0 0(1) (4) A. otiosus Say males2 1 0 0 0 1 A. otiosus-relative females2 (205) (106) (193) (2) (506) A. planipennis Fairmaire 22(26) 48(46) 20(44) 31(59) 121(175) A. ruficollis (Fabricius) 1(0) 0 1(1) 0 2(1) A. transimpressus Fall males2 2 1 0 1 4 A. vittaticollis (Randall) 0(1) 0 0 0 0(1) UnidentifiedAgrilus 0(19) 0(5) 0(2) 0(0) 0(26)

Anthaxia viridicornis (Say) 2(0) 0 0 0 2(0) An. viridifrons Gory 1(3) 0 0 0 1(3) UnidentifiedAnthaxia females (0) (0) (1) (0) (1)

Chrysobothris femorata (Olivier) 0 0 0(2) 0 0(2) C. quadriimpressa Gory&LaPorte 0 0(1) 0 0 0(1)

Dicerca lurida (Fabricius) (0) (1) (0) (0) 0(1)

Total buprestids 104(287) 79(169) 85(264) 33(66) 301(786)

1Females of these two species cannot be distinguished from one another. 2In the A. otiosus species group or a close relative. Females in this group cannot be distinguished from one another. See MacRae (2003) for a complete list of species in this group. 26 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 or Agrilus captured did not vary significantly among the trap colors (Fig. 5a, b). Also, the mean number of buprestid or Agrilus species captured did not vary significantly among trap colors (Fig. 6a, b). Considering the more common Agrilus species captured, the mean percentage of EAB captured per trap did not vary significantly among the different trap colors (Fig. 5c). The mean percentage of A. otiosus-relatives that were males, females, or for both sexes combined per trap was significantly higher on green traps compared to purple traps, while the percentage captured on green over purple or purple over green combination traps tended to be intermediate (Fig. 5d). Four species of male A. otiosus-relatives were captured, of which 96% were A. masculinus (Table 2). The mean percentage of A. celti + A. egenus or Agrilus lecontei Saunders captured per trap did not vary significantly among trap colors (Fig. 5e). However, the percentage of A. lecontei females captured was almost significant P( = .0555) with more tending to be captured on green traps than on purple traps (Fig. 5f).

Discussion Some studies have confirmed that visual cues are used by several species of Buprestidae to locate and select their mates (Gwynne and Rentz 1983; Lelito et al. 2008, 2011; Domingue et al. 2011), so it is likely that visual cues are also important for buprestids to locate and select their host plants. The host range of buprestids includes most woody species (both conifers and hardwoods) but most buprestids are restricted to hosts within a single plant family or genus (Nelson et al. 2008). Given the variability in color and shape among buprestid adults and the variation in hue and reflectance among tree species (Campbell and Borden 2005), it is not surprising that trap preference varied among the buprestid genera and species collected in our study. In the trap-shape study, purple Agrilus silhouette traps captured the most buprestid and Chrysobothris species, while green Agrilus silhouette traps captured the most Agrilus species. The Agrilus shape combined with the green or purple color of the silhouette may explain why these traps were most attractive. Also, the white background surrounding the silhouettes may have enhanced attraction to these traps. Background contrast is believed to play an important role in host location by insects (Prokopy and Owens 1983). We realize that the treatments used in the present study limit the strength of interpretation that can be made regarding the effects of the white background and the Agrilus silhouettes. Nevertheless, in our trap color study where no white background colors were used, significant differences were not detected for the number of buprestid or Agrilus species captured among green, purple, or green-purple combination traps. Future studies are warranted to compare different background colors with Agrilus silhouettes and other shapes to further elucidate differences in attractiveness of these types of traps. Although, trap color or shape did not significantly affect the overall number of buprestid individuals captured, genus and species level differences were found. EAB decoys and Agrilus silhouettes, with the exception of the enlarged EAB photograph, tended to increase attraction of EAB to the traps. Although the EAB photograph appeared to the human eye as the same color as an actual EAB adult beetle, it was not very attractive to EAB in our study. It is possible that EAB perceive color and light reflecting from two-dimensional photographs differently than they would from actual three-dimensional insects. Furthermore, the cuticle of EAB adults is iridescent and reflects many colors due to pigmen- tation and structural characteristics (Baker et al. 2011), which may not be the same as a two-dimensional photograph. The effect that the insect trapping glue may have had on the distortion and reflectance of the EAB photograph should also be acknowledged. Crook et al. (2009) found that the insect trapping glue used in their study, which was very similar in transparency and consistency 2013 THE GREAT LAKES ENTOMOLOGIST 27 to the glue we used, increased reflectance by 2.5%. It is also possible that the white background that surrounded the EAB photograph deterred attraction, however, the white backgrounds of the purple or green Agrilus silhouette traps did not appear to reduce attraction of EAB. We expected attraction to traps with EAB decoys to be higher for male EAB compared to females, given that Agrilus males usually seek females for mating rather than vice versa. Lelito et al. (2008) found only males to alight on ash leaflets where dead EAB adults had been placed. Therefore, it is puzzling that the addition of 25 EAB decoys to the purple rectangular traps in our study increased the overall attraction of EAB females, but not males. Francese et al. (2010) found EAB females to be more attracted to purple than males, and suggested that purple mimics the color of tree bark to EAB. It is possible that EAB females may orient toward other EAB adults, especially if they are against certain background colors such as purple because their presence indicates the location of a suitable host, i.e., females may tend to land on a host near other EAB adults. Lelito et al. (2007) observed wild EAB males in the field that hovered above dead beetles pinned to the ash leaflets and then rapidly and accurately dove onto the body of the beetle decoy. Therefore, it is possible in our study that some males landed on the dead EAB decoys and were able to fly away without being captured because the decoys were not coated with Pestick™. Furthermore, male EAB are more attracted to green compared to females (Francese et al. 2010), and perhaps, if we had placed EAB decoys on a green background in our study the response of males to the EAB decoys would have been stronger. Responses by A. anxius to the various trap designs and colors were different from those of EAB. Significantly more A. anxius males were captured on the enlarged EAB photograph traps than on purple rectangle traps with or without EAB decoys, and captures on green and purple Agrilus silhouette traps were intermediate. Although significant differences were not detected,A. anxius females followed a similar pattern as the A. anxius males. It is unclear whether the silhouettes themselves attracted A. anxius males or if attraction was due primarily to the white background that surrounded the EAB photographs and the purple and green Agrilus silhouettes. Supporting this latter hypothesis is the fact that the background bark color of many birch (Betula) species is white, which is likely an important visual cue for this Agrilus species to locate its host. Agrilus anxius was also found to be attracted to white corrugated plastic sticky traps baited with host volatiles (Gary Grant, Canadian Forest Service, personal communication 2009, now deceased). While significant differences in the percentage of EAB adults captured per trap were not detected among purple, green and purple-green combination traps, there was a general trend of male EAB to be more attracted to green, while female EAB were more attracted to purple. This trend is consistent with the findings of previous studies by Crook and Mastro (2010) and Francese et al. (2010). Green is apparently much more attractive than purple for the A. otiosus- relatives. Both sexes were most attracted to green-colored traps in both of our studies. Attraction to green may be host related for the A. otiosus-relatives because members of this group tend to oviposit on small branches (Hespenheide 1969, MacRae 1991) that would most often be in the canopy where green-colored foliage is typically present. Furthermore, most A. otiosus-relatives attack dead or severely weakened branches rather than healthy live branches (MacRae 1991), and the shade of light green used for our traps may appear similar to foliage of weakened or dying trees, i.e., chlorotic foliage that is slightly yellow or lighter in color. Although significant differences were not detected, Agrilus lecontei and A. egenus + A. celti responded in a similar pattern in our trap color study and tended to prefer green colored traps. 28 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

In the trap shape study, Chrysobothris species were most attracted to the purple Agrilus silhouette traps. Interestingly, the green Agrilus silhouette traps were among the least attractive traps for Chrysobothris. Furthermore, no Chrysobothris were captured on the all-green traps in our trap color study, albeit, only three Chrysobothris individuals were captured in that study. Oliver et al. (2003) found colors in the red spectrum (which would include purple) to be most attractive to buprestids, especially Chrysobothris. Moreover, Oliver et al. (2003) found white traps were moderately attractive to Chrysobothris spp., so perhaps it was the combination of the white and purple that enhanced attraction of Chrysobothris in our trap shape study. Future studies that include traps with different purple shapes and all white traps would help determine if the purple Agrilus silhouette contributed to the attraction of Chrysobothris to this type of trap or if it was merely attraction to the two colors combined. In the present study, attraction to different trap colors and shapes varied among buprestid genera and species. Therefore, to effectively monitor buprestid populations, multiple trap shapes and colors should be considered when conducting general surveys. More studies are warranted to determine the most attractive color and shape combination for individual buprestid species, especially those of economic importance.

Acknowledgments We thank Kyle Brown, Stephen Burr, Tina Ciaramitaro and Alison Barc for field and laboratory assistance; Stan Wellso and Jason Hansen for buprestid identifications; Joseph Francese, Jason Hansen and two anonymous reviewers for comments on a previous draft of this manuscript.

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Abstract We assessed the physicochemical and biological continuity of a 2nd–4th order reach of the Little Manistee River in northern Lower Michigan. Contrary to typical woodland streams, the downstream sites of the river were covered with a dense riparian canopy, whereas the upstream sites were devoid of this canopy due to historical (≥10 ybp) agriculture. Other than slight changes in water temperature and dissolved oxygen, there were no appreciable differences in measured water physicochemistry between the canopied and non-canopied sites. The stream, however, appeared biologically discontinuous as indicated by lower shredder abundance and higher filtering collector abundance at the upstream (non-canopied) sites for both benthic macroinvertebrate and adult caddisfly assemblages. Benthic scraper abundance was, likewise, higher in the upstream sites. Our results suggest that changes in riparian canopy alone can lead to changes in biological assemblages, even without obvious changes in water physicochemistry.

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The river continuum concept (Vannote et al. 1980) describes a predictable pattern of gradual changes in river morphometrics and organismal assemblages as width increases and the river interacts with its riparian corridor. Anthro- pogenic disturbances, however, may disrupt this continuum, particularly in small–medium streams (Pringle et al. 1993; Delong and Brusven 1993, 1998; Houghton 2006, 2007). Removal of the riparian canopy cover, for example, may cause a loss of coarse allochthonous input needed by invertebrate shredders (Sabater et al. 2000, Warren et al. 2007). Agricultural streams in particular have concentrations of sediment, nutrient, and fine organic matter input hun- dreds of times higher than undisturbed streams (Royer et al. 2004, Inwood at al. 2005). Riparian habitat loss, with subsequent increases in nutrient and sediment input, has been repeatedly found to be the most widespread stressor of streams in the U.S. generally, in the Plains and Lowlands region of the U.S., and in Michigan (Paulsen at al. 2008, Wang et al. 2008). It is difficult to separate the habitat loss and sediment input components of riparian disturbance, as they frequently occur together. Several recent studies of agricultural watersheds, however, have noted positive correlations between the amount of intact riparian vegetation and the biological diversity in the adjacent streams (Houghton 2004a, Rios and Baily 2006, Urban et al. 2006). More specifically, Houghton et al. (2011a) found that sites in a southern Michigan agricultural stream protected by riparian canopy had 3× the number of adult caddisfly species as sites without canopy, even though physicochemistry remained unchanged and typical of agricultural streams. These studies suggest the hypothesis that both riparian habitat loss and increases in sediment

3Department of Biology, 33 East College Street, Hillsdale College, Hillsdale, MI 49242. (e-mail: [email protected]). 32 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 input are individually important for affecting stream biota. The purpose of our study, therefore, was to test this hypothesis by observing the effects of riparian canopy loss on stream macroinvertebrate assemblages in a stream without obvious sediment input.

Materials and Methods Study site. The Little Manistee River is located in the northwestern portion of the Lower Peninsula of Michigan. It is approximately 100 km in length and drains a watershed of approximately 1100 km2 before draining into Lake Michigan. Over half of the watershed is officially protected by the Manistee National Forest and the Pere Marquette State Forest, and much of the remaining land is also undeveloped (Fig. 1). Due to its relatively undisturbed watershed and stable groundwater input, it is one of the coldest and most stable streams in the Lower Peninsula (Tonello 2005). It hosts potomadromous spawning runs of steelhead [Oncorhynchus mykiss (Walbaum)], Chinook salmon [Oncorhynchus tshawytscha (Walbaum)], and brown trout [Salmo trutta Linnaeus] from Lake Michigan, and contains breeding populations of brook trout [Salvelinus fontinalis (Mitchell)] (Seelbach 1993). Although much of the watershed is protected, the headwaters of the Little Manistee are primarily composed of pasture and feral fields (Tonello 2005). From our observations, it has been ~10 years since the upper watershed was under active cultivation. Five sites were sampled on the Little Manistee during this study. Sites 1–2 were located in the non-canopied headwaters region and sites 3–5 were all within the forested State and National Forest land (Fig. 1). All 5 sites were

Figure. 1. Aerial photograph (Google Earth) of the study area showing the location of the Little Manistee River, its important tributaries, and the location and photographs of our five collecting sites. Green areas denote forest, whereas lighter brown areas denote agriculture, feral fields, or urban landuse. 2013 THE GREAT LAKES ENTOMOLOGIST 33 within a stream reach of 10 km. All sites were upstream from major tributaries. Stream width ranged from 3 m (2nd order) in the upper sites to 10-12 m (4th order) at the lower sites. Determination of the extent of canopy cover was from aerial maps and visual inspection. Except for the occasional alder (Alnus sp.) along the banks, non-canopied areas of both streams were almost devoid of plants > 2 m in height within 100 m of either bank. Thus, the distinction between ‘canopied’ and ‘non-canopied’ sites was distinct enough that further quantification and precision were deemed unnecessary. Physicochemical sampling. Physicochemical measurements were made twice during June and once monthly from July through September 2011. Six sets of measurements were made from each site on each date. Conductiv- ity (ECTestr Low, Eutech Instruments), pH (AccuMet AP61, Fisher Scientific), dissolved oxygen (YSI-55, YSI Environmental), and temperature (YSI-55, YSI Environmental) measurements were all made on-site. All measurements were made within 2 h of each other to minimize diel fluctuations. Benthic macroinvertebrate sampling. Benthic macroinvertebrates were sampled using Hess samplers with 0.3 m2 areas (Barbour et al. 1999). Six Hess samples were taken from each site of each stream within a diversity of habitats. Sites 1, 2, and 4 were sampled during May 2010 and sites 3 and 5 were sampled in May 2011. All benthic specimens were identified to the lowest identifiable taxon, typically genus, after Hilsenhoff (1995). Adult caddisfly sampling. Adult caddisflies were sampled using ultraviolet light traps, which consisted of an 8-watt ultraviolet light placed over a white pan filled with 70% ethanol. Each trap was placed within 2 m of a sampling site at dusk and retrieved approximately two hours later. By stan- dardizing the time of collection, wattage of the light source, and size of collecting pan, the technique yielded quantitative samples of the nocturnally active caddisfly adults and allowed for comparisons between sites (Houghton 2004a). To standardize weather conditions, samples were collected only if the peak daytime temperature was > 22°C, dusk temperature was >13°C, and there was no noticeable wind or precipitation at dusk. Sampling occurred approximately bi-weekly during June and July 2011, the peak emergence period of caddisflies in northern Michigan (Houghton et al. 2011b), for a total of 5 samples from each site. All adult specimens were identified to species after Houghton (2012), except for females of Hydroptilidae, Hydropsychidae, and Polycentropodidae, which lack the characters necessary for doing so. Such specimens were not included in any analyses. Data analyses. Specimens of both benthic invertebrates and adult caddisflies were placed into trophic functional groups following Merritt et al. (2008). Algal piercers were considered gathering collectors in analyses. Mean percentages of the functional groups informative of stream condition at each site: scrapers, shredders, and filtering collectors (e.g., Allan 1995, Houghton 2007), were compared to each other by a one-way Analysis of Variance. Percentages were transformed through an ArcSine function before analysis (Zar 2007). Mean water physicochemical values were compared using a 2-way Analysis of Vari- ance to determine differences between both sampling site and sampling day. Sampling sites were examined for patterns in their benthic macroinvertebrate and adult caddisfly assemblages with Detrended Correspondence Analysis (DCA) using the program PC-ORD for Windows® (McCune and Grace 2002). The DCA analysis was performed on a two-dimensional data matrix of sampling sites by taxa relative abundance values. Relative abundances were determined by counting the number of specimens collected at each site and then coding 0 specimens as ‘0’, 1–10 as ‘1’, 11–100 as ‘2’, 101–1000 as ‘3’, and 1001–10,000 as ‘4’. Since data coding accounted for variation in specimen abundance between sites, it was a more powerful measure than simple presence or absence data

(Feminella 2000, Houghton 2004a). Coding on a log10 scale mitigated the effects 34 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 2. Detrended correspondence analysis ordination of (A) adult caddisflies, (B) benthic invertebrate samples from the Little Manistee River.

of outlier samples often associated with light-trapping data, as well as the influence of highly abundant species (Cao et al. 1997, Houghton 2004a, Houghton et al. 2011b). All taxa were weighted equally in each analysis.

Results Assemblages of adult caddisflies and benthic invertebrates were different between sites with low riparian canopy and sites with higher canopy (Fig. 2); sites 1–2 appeared distinct from sites 3–5. Adult caddisfly filtering collector abundance was higher at sites 1–2 and lower at sites 3–5, although there were indistinct groups among the latter sites (Fig. 3). Shredder abundance was higher at sites 3–5, and lower at sites 1–2. Scraper abundance was unchanged between sites. The combined abundance of shredder, scraper, and filtering collectors was > 80% of total adult caddisfly specimens. Benthic invertebrates exhibited the same trends in shredder and filtering collector abundances: the former was more abundant at sites 3–5 and the latter was more abundant at sites 1–2 (Fig. 3). Scraper abundance was higher at sites 1–2 and lower at sites 3–5. All sites were dominated by gathering collectors (60–80%), and the combined abundance of shredder, scraper, and filtering collectors was ≤ 30% of total specimens at all sites (Fig. 3). Individual filtering collector species were most abundant at sites 1–2, whereas shredder species were at their highest abundance at sites 3–5 (Table 1). The most abundant filtering collectors at the non-canopied sites were Brachycentrus americanus (Banks) (Brachycentridae), Cheumatopsyche oxa Ross, Hydropsyche slossonae Walker, and H. sparna Ross (Hydropsychidae). Hydropsyche sparna (Hydropsychidae), B. americanus, and Lepidostoma togatum Hagen were among the top 10 at all 5 sites. Conductivity and pH exhibited no significant differences between sampling sites or between sampling dates (Fig. 4). Sites 1 and 2 had higher temperatures and lower dissolved oxygen than the other sites. Although there was significance between the temperature and dissolved oxygen values of different sampling dates in the overall model, there was also considerable overlap between these dates. 2013 THE GREAT LAKES ENTOMOLOGIST 35

Figure 3. Mean (+SE) values of percentage of filtering collectors, scrapers, and shredders for benthic larvae and adult caddisflies of the Little Manistee River. Bars topped with the same lowercase letter were not statistically different (1-way Analysis of Vari- ance with post-hoc Tukey test). n.s. = not significant. 36 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 4. Mean physicochemical measurements determined on 5 different days from multiple sites of the Little Manistee River, with associated P-values (two-way Analysis of Variance). n.s. = not significant. Asterisks signify significantly different means -be tween sites; one-way Analysis of Variance with (post-hoc Tukey test). (A) pH, (B) water temperature, (C) conductivity, (D) dissolved oxygen. Conductivity and pH displayed on expanded scale to show detail and with connecting lines omitted for clarity. Error bars omitted for clarity.

Discussion The lack of riparian canopy at the upstream sites does not appear to have obvious effects on measured water physicochemistry. Conductivity and pH collectively have been found to explain nearly 80% of the watershed disturbance variation between sites of New Jersey watersheds, and were also associated with differences in organismal assemblages (Zampella and Laidig 1997, Dow and Zampella 2000). Conductivity, in particular, is often used as a preliminary indicator of nutrient, sediment, and organic matter concentrations. Such concentrations accumulate naturally in larger rivers, or anthropogenically in disturbed streams (Allan 2004). Michigan streams disturbed by agriculture have levels > 2× that of the Little Manistee (Castillo et al. 2000, Houghton et al. 2011a). Our low conductivity levels, as well as the lack of difference in both variables between study sites, suggested no important differences in natural or anthropogenic sediment input throughout the continuum. It is difficult to judge the importance of temperature and dissolved oxygen differences between sites due to the differences in both variables between sampling dates. The temperature difference between sampling dates was similar (~5° C) to 2013 THE GREAT LAKES ENTOMOLOGIST 37

Table 1. The 10 most abundant caddisfly species from 5 sites of the Little Manistee River.

Species Functional group Percentage of fauna

Site 1 1 Hydropsyche sparna Ross Filtering collector 31% 2 Hydropsyche slossonae Banks Filtering collector 20% 3 Cheumatopsyche oxa Ross Filtering collector 7% 4 Lepidostoma togatum (Hagen) Shredder 6% 5 Brachycentrus americanus (Banks) Filtering collector 4% 6 Neureclipsis crepuscularis (Walker) Filtering collector 3% 7 Lype diversa (Banks) Gathering collector 3% 8 Psychomyia flavida Hagen Gathering collector 2% 9 Glossosoma nigrior Banks Scraper 2% 10 Hydroptila consimilis Morton Gathering collector 2% Site 2 1 Hydropsyche sparna (Ross) Filtering collector 35% 2 Cheumatopsyche oxa Ross Filtering collector 18% 3 Cheumatopsyche gracilis (Banks) Filtering collector 8% 4 Hydropsyche slossonae (Banks) Filtering collector 6% 5 Neureclipsis crepuscularis (Walker) Filtering collector 5% 6 Psychomyia flavidaHagen Gathering collector 5% 7 Glossosoma nigrior Banks Scraper 4% 8 Hydroptila consimilis Morton Gathering collector 4% 9 Lepidostoma togatum (Hagen) Shredder 4% 10 Brachycentrus americanus (Banks) Filtering collector 3% Site 3 1 Lepidostoma togatum (Hagen) Shredder 30% 2 Lype diversa (Banks) Scraper 22% 3 Hydropsyche sparna (Ross) Filtering collector 10% 4 Brachycentrus americanus (Banks) Filtering collector 7% 5 Lepidostoma bryanti (Banks) Shredder 6% 6 Neureclipsis crepuscularis (Walker) Filtering collector 5% 7 Glossosoma nigrior Banks Scraper 4% 8 Nyctiophylax affinis (Banks) Predator 4% 9 Hydroptila consimilis Morton Gathering collector 3% 10 Hydroptila jackmanni Blickle & Morse Scraper 2% Site 4 1 Lepidostoma togatum (Hagen) Shredder 26% 2 Lepidostoma bryanti (Banks) Shredder 11% 3 Brachycentrus americanus (Banks) Filtering collector 9% 4 Lype diversa (Banks) Scraper 8% 5 Glossosoma nigrior Banks Scraper 6% 6 Neureclipsis crepuscularis (Walker) Filtering collector 5% 7 Nyctiophylax affinis(Banks) Predator 5% 8 Hydroptila jackmanni Blickle & Morse Scraper 3% 9 Cheumatopsyche gracilis (Banks) Filtering collector 3% 10 Hydropsyche sparna (Ross) Filtering collector 2% Site 5 1 Lepidostoma togatum (Hagen) Shredder 25% 2 Brachycentrus americanus (Banks) Filtering collector 16% 3 Lepidostoma bryanti (Banks) Shredder 14% 4 Psychomyia flavidaHagen Gathering collector 7% 5 Helicopsyche borealis (Hagen) Scraper 6% 6 Ceraclea transversa (Hagen) Gathering collector 5% 7 Hydropsyche sparna (Ross) Filtering collector 5% 8 Cheumatopsyche oxa Ross Filtering collector 4% 9 Lype diversa (Banks) Gathering collector 4% 10 Polycentropus pentus Ross Predator 4% 38 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 that between sampling sites. The observed differences between sites were likely due to the lack of canopy cover in the headwaters allowing additional sunlight to reach the stream surface and warm the water. Organisms at the canopied and non-canopied sites were probably exposed to slightly different temperature profiles. Dissolved oxygen typically exhibits a reciprocal relationship with temperature; thus, it was lower where temperature was higher in both streams (Allan 1995). While the physicochemical differences between sites were unclear, our observed organismal assemblages suggested a reversal of predicted biological continuity. Continuity theory (Vannote et al. 1980) predicts high shredder abundance in low order (upstream) stream sites due to the high relative abundance of riparian canopy cover. Instead, Little Manistee shredders were more abundant at the higher order (downstream) sites. We suspect that this observation was due to canopy cover actually increasing in the downstream sites relative to the upstream sites, even though the Little Manistee obviously widens downstream (Fig. 1). Scrapers typically increase in abundance into the 4–5th order, and filtering collectors reach their highest abundances at the highest stream orders. In the Little Manistee, both functional groups instead decreased at the downstream sites. While there have been many critiques and modifications of stream continuity theory (most recently Thorp et al. 2006), small woodland streams, such as the Little Manistee, do tend to follow the general predicted patterns if they are undisturbed (Allan 2004). Conversely, disturbed small woodland streams do not follow predicted patterns. Instead, they typically exhibit abundant filtering collectors and few shredders (Pringle et al. 1993; Delong and Brusven 1998; Houghton 2006, 2007). The abundance of filtering collectors alone accounted for nearly 70% of the watershed disturbance variation of small and medium streams in Minnesota (Houghton 2006). The overall assemblages of the upstream Little Manistee sites were those predicted from a medium-large river, despite a width of 3–4 m. More specifically, the relative abundances of adult caddisfly shredders and filtering collectors corresponded to those of 20–30 m wide Minnesota rivers (Houghton 2007). Once canopy cover returned downstream, however, continuity normalized and was maintained for the remainder of the studied stream length. Even though filtering collectors dominated the non-canopied sites, these assemblages did not indicate ecosystems disturbed by excess sediment or nutrient input. Typically, a small stream with high levels of anthropogenic disturbance has an abundance of the specific filtering collector species that are normally found in large rivers. These species increase in disturbed small streams due to an abundance of fine particulate organic matter that they can utilize as a food source (Allan 1995, Barbour et al. 1999, Allan 2004, Houghton 2007). For example, the hydropsychid species Cheumatopsyche campyla Ross, Hydropsyche simulans Ross, and Potamyia flava Hagen are all typically found in large rivers of Minnesota, and only rarely in undisturbed small streams (Houghton 2012). In small agricultural streams, however, these same species were abundant and constituted significant indicators of disturbance statewide (Houghton 2004b). In the Little Manistee, all of the abundant filtering collectors (Table 1) were those naturally found in small woodland streams of the northcentral U.S. (Houghton et al. 2011b, Houghton 2012), albeit at lower relative abundances. Thus, instead of an increase in small-particle filtering collectors associated with large rivers and polluted small streams, our sites suggested a shift in the relative abundances of invertebrate assemblages already present. The normalization of continuity within ~3 km after canopy returned (between sites 2 and 3) also suggested the local effect of small-scale canopy loss, rather than systemic disturbance, such as sediment input or canopy removal on a large scale. Our results also suggest relative strengths and weaknesses of using benthic macroinvertebrates and adult caddisflies to assess stream conditions. In the case of the former, nearly all of our sites were dominated by gathering collectors, which are common in nearly all habitat types and usually not informative of stream conditions (Vannote et al. 1980, Stagliano and Whiles 2002, 2013 THE GREAT LAKES ENTOMOLOGIST 39 Houghton 2007, Whiting et al. 2011). Thus, the high abundance of non-informative taxa relative to the informative scrapers, shredders, and filtering collectors may decrease the ‘signal’ relative to the ‘noise’ of a sample and render determination of stream conditions more difficult (e.g., Cao and Hawkins 2011). In the case of the latter, the lack of response from the scraper functional group has been noted in previous studies (Houghton 2006, 2007). While scraper taxonomic richness may be proportional within the Trichoptera (e.g., Wiggins 1996), scraper specimens are not generally abundant in blacklight samples relative to filtering collectors or shredders, or in benthic samples compared to taxa of other orders (Houghton 2012). Thus, the metric may not be abundant enough to be informative of stream conditions when using the Trichoptera exclusively. Both of our assemblages reinforce the assertion that a loss of riparian habitat from ≥ 10 years ago can still impact stream biota (Harding et al. 1998). The potential mechanisms of this impact—such as decrease in allochthonous CPOM, increase in allochthonous FPOM, changes in substrate composition, increase in flooding, or other physiochemical factors—still need to be worked out. Year-round data logging would be beneficial to determine subtle difference in temperature between sites, as would more direct measurements of nutrients or suspended organic matter.

Acknowledgments Research costs were paid by the Mossberg Conservation Fund and Hillsdale College biology department funds. We thank Kelsey Brakel for field assistance and Mark Nussbaum for water physicochemical assistance. The valuable comments of Matt Whiles and Rebecca Shealy improved earlier versions of the manuscript. This is paper #5 of the G.H. Gordon BioStation Research Series.

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Seelbach, P. W. 1993. Population biology of steelhead in a stable-flow, low gradient tributary to Lake Michigan. Transactions of the American Fisheries Society 122: 179-198. Stagliano, D. M., and M. R. Whiles. 2002. Macroinvertebrate production in a prairie stream. Journal of the North American Benthological Society 21: 97–113. Thorp, J. H., M. C. Thoms, and M. D. Delong. 2006. The riverine ecosystem synthesis: biocomplexity in river networks across space and time. River Research and Applications 22: 123–147. Tonello, M. A. 2005. Little Manistee River. Michigan Department of Natural Resources Status of the Fisher Resource Report. Available from http://www.mich.gov/documents/2005-8_Little-Manistee_River_144067_7.pdf (accessed 09 March 2011). Urban, M. C., D. K. Skelly, D. Burchsted, W. Price, and S. Lowry. 2006. Stream communities across a rural–urban landscape gradient. Diversity and Distributions 12: 337–350. Vannote, R. L., G. W. Minshall, K. W. Cummins, J. R. Sedell, and C. E. Cushing. 1980. The river continuum concept. Canadian Journal of Fisheries and Aquatic Sciences 37: 130–137. Wang, L., T. Brenden, P. Seelbach, A. Cooper, D. Allan, R. Clark, and M. Wiley. 2008. Landscape based identification of human disturbance gradients and reference conditions for Michigan streams. Environmental Monitoring and Assessment 141: 1–17. Warren, D. R., E. S. Bernhardt, R. O. Hall, G. E. Likes. 2007. Forest age, wood, and nutrient dynamics in headwater streams of the Hubbard Brook Experimental Forest, NH. Earth Surface Processes and Landforms 32: 1154–1163. Whiting, D. P., M. R. Whiles, and M. L. Stone. 2011. Patterns of macroinvertebrate production, trophic structure, and energy flow along a tallgrass prairie stream continuum. Limnology and Oceanography 56: 887–898. Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera), nd2 edition. University of Toronto Press, Ontario. Zampella, R. A., and K. J. Laidig. 1997. Effect of watershed disturbance on pinelands stream vegetation. Journal of the Torrey Botanical Society 124: 52–66. Zar, J. H. 2007. Biostatistical Analysis, 5th edition. Prentice-Hall, Englewood Cliffs, NJ. 42 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Diversity and Community Structure of Stream Insects in a Minimally Disturbed Forested Watershed in Southern Illinois J. E. McPherson1, Jacqueline M. Turner1,3, and Matt R. Whiles1,2

Abstract The Lusk Creek Watershed, located in Pope County, IL, long has been rec- ognized as a high quality area of biological significance, but surveys of the stream macroinvertebrate fauna have been limited. Thus, a survey of the benthic insect community at 11 sites in the upper portion of Lusk Creek was conducted from May 2003 to April 2005. A total of 20,888 specimens, mostly immatures, were examined during the study and represented eight orders. The Diptera, by far, was the most abundant order, with 18,590 specimens, almost all of which were members of the Chironomidae or Simuliidae. Members of the EPT (Ephemer- optera, Plecoptera, Trichoptera) contributed 1,550 specimens. The Coleoptera was represented by 647 specimens, most of which were members of Stenelmis (Elmidae) (n = 612). The Shannon diversity index (H´) ranged from 1.07-2.01 for individual sites and was indicative of relatively undisturbed streams in this region. Jackknife analyses of richness estimated that as many as 37 taxa were unobserved in this survey. Results provide information on reference conditions in the region and a foundation for future monitoring.

______

Illinois, known unofficially as the prairie state (Shankle 1938), has under- gone major landscape changes during the last 200 years, primarily because of agricultural and industrial development. Not unexpectedly, these anthropogenic changes have been concentrated in the highly populated regions of northern Illinois (IDENR 1994a) and the heavily farmed areas of central Illinois (IDENR 1994b). Prior to intensive European settlement, over half of Illinois was prairie (8,741,209 ha) with the remainder primarily forest (5,584,661 ha) that often was concentrated along rivers (Anderson 1970, Iverson et al. 1989). By 1924, the total forest acreage of Illinois had been reduced to just over 1,214,056 ha (Telford 1926). Klopatek et al. (1979) estimated that by 1967, only 11% of Il- linois’ natural vegetation still remained. Inherent with land use changes since settlement, most Illinois streams have been severely altered, with major manipulations including channelization and construction of impoundments. Channelization has occurred primarily for agricultural practices (IDENR 1994b), with 22.7% of streams in the state affected (Mattingly and Herricks 199l). Impoundments have been constructed for transportation, flood control, water supplies, and recreation (IDENR 1994b). Southern Illinois also has experienced anthropogenic changes but to a much lesser extent than the central and northern regions of the state. In 1820, the southern portion of Illinois was covered almost completely by forest (Anderson 1970, Iverson et al. 1989). At that time, Pope County had 95,627 ha of forest (Iverson et al. 1989) and a human population of 2,610 (Pope County Historical Society 1986). By 1890, the population had reached its peak of 14,017

1Department of Zoology, Southern Illinois University, Carbondale, IL 62901. 2Center for Ecology, Southern Illinois University, Carbondale, IL 62901. 3Current Address: 198 Mt. Zion Road, Crawfordville, FL 32327. 2013 THE GREAT LAKES ENTOMOLOGIST 43 (Pope County Historical Society 1986). By the early 1900s, most of the forest cover of the Shawnee Hills region in southern Illinois had been removed, and most land was being farmed intensively (Kandl 1990). By 1924, the forested area of Pope County, which includes much of the Shawnee Hills, had been reduced to 26,409 ha (Telford 1926). By the beginning of the Depression, the land was no longer profitable for farming and much of it had been “forfeited” (i.e., foreclosed). As a result, the eastern Shawnee Hills region was selected as the site of a national forest (Soady 1965). By 1980, the population had decreased to 4,250 (Pope County Historical Society 1986). By 1985, forest acreage in the county had increased to 60,379 ha (Iverson et al. 1989). Baseline studies and reference conditions. Studies of anthropogenic impacts, both biological and physical, ideally would include baseline studies preceding human influence for comparison. As that generally is not possible, baseline studies of systems with limited human influence often are used to es- tablish reference points for assessing ecological integrity (Metzeling et al. 2006). White (1978) surveyed Illinois natural areas and found, based on 1,089 sites, only 10,409 ha of high quality natural communities. Large areas of the state had no high quality sites, including the former central prairies that had been almost completely farmed; other areas had clustered sites that were located near the western and southern borders of Illinois along rivers and bluffs. The 11 southernmost counties had 396 sites, including Pope County with 88 sites, of high quality natural communities (White 1978); T. G. Kieninger (personal communication) stated that the 396 and 88 sites included 2,134 ha and 129 ha, respectively. This high number of sites and large amount of high quality acreage in southern Illinois reflects the lower degree of anthropogenic impacts in this region. Lusk Creek. The Lusk Creek watershed lies entirely within Pope County (Figs. 1A and B). Draining north to south, it is ca. 40-km in length. It has high topographic relief (Fig. 2), which apparently has protected the major stream valleys from human influence (Hudak 1979). This watershed has received several designations and ratings that recognize the high quality of the area (INPC 2008, NPS 2009, Hite and Bertrand 1989). In 1990, a portion of Lusk Creek, including the adjacent watershed (from the Lusk Creek/Dog Hollow confluence to Ramsey Branch [USFS 1996]), was designated a National Wilderness Area, which is a federal land designation where natural processes dominate the landscape and the human presence is “substantially unnoticeable” (United States Congress 1990). In 1992, Lusk Creek from Flick Branch to Quarrel Creek and from Manson Ford to Little Lusk Creek and Copperous Branch were rated as “Biologically Significant Streams” (Page et al. 1992). In 1993, Lusk Creek from Copperous Branch to just upstream of Ramsey Branch was rated as a “Unique Aquatic Resource” (Class A Stream), and the remaining upstream and downstream portions of Lusk Creek and the entire length of Little Lusk Creek were rated, collectively, as a “Highly-valued Aquatic Resource” (Class B Stream) (Bertrand et al. 1996). Changes in diversity of stream fauna. The Nature Conservancy and others in the 1980s and 1990s evaluated the conservation status of over 30,000 terrestrial and aquatic species and subspecies with existing state natural heritage database information. Fourteen groups of plants and animals, representing 20,900 species, had enough associated information to evaluate the entire taxonomic assemblage (Master et al. 2000). Of the 14, four groups of freshwater animals comprised the highest proportion of species at risk (i.e., presumed/possibly extinct, critically imperiled, imperiled, and vulnerable). These included mussels, crayfish, stoneflies, and fish with 69%, 51%, 43%, and 37% at risk, respectively. Not enough data were available to evaluate the conservation status of mayflies or caddisflies (Master et al. 2000). As with the national trends, the diversity of Illinois stream fauna also has declined as evidenced by decreases in the same four groups noted above. Of 44 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Fig. 1. Locations of Lusk Creek and watershed in Pope County, Illinois. (A) Watershed. (B) Lusk Creek. 2013 THE GREAT LAKES ENTOMOLOGIST 45

Fig. 2. Lusk Creek watershed within Shawnee Hills Illinois Natural Division (SH), which is subdivided into Greater SH (GSH) and Lesser SH (LSH). Shown here is range of elevations (GSH, 104–256 m, LSH, 98–207 m). 46 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

79 mussel species in the state, 17 (22%) are extinct or extirpated (INHS 2012a) and 25 (32%) are endangered or threatened (Mankowski 2010, INHS 2012a). Of the 22 crayfish species, one (5%) has disappeared (IDENR 1994b) and four (18%) are endangered (IDENR 1994b, Mankowski 2010). Of the 77 stonefly species, 22 (29%) are extinct or extirpated, including two Illinois endemics, and 19 (25%) are “critically imperiled” (DeWalt et al. 2005). Of the 187 original fish species, 11 (6%) species have disappeared (IDENR 1994b) and 31 (17%) are threatened or endangered (Mankowski 2010). Smith (1971) attributed the richness of Illinois fishes to the high number of streams and the associated diversity of habitats. However, he identified seven factors that have reduced Illinois fish diversity. They include siltation, drainage of lakes and wetlands, dessication during drought, species interactions (due to habitat modification and species introduction), pollution, impoundments, and temperature disruption. Page (1991) agreed with Smith (1971) and suggested that these same factors were major threats to stream biodiversity in general. Stone et al. (2005) investigated streams draining agricultural landscapes in southwestern Illinois. They found that the benthic invertebrate communities in these systems were dominated by pollution-tolerant taxa and identified siltation as the major human impact affecting biological integrity. They also found that the insect portion of the community increased as riparian buffers increased in width. Heatherly et al. (2007) investigated streams across Illinois that represented the major natural divisions and a land-use continuum. They found that physical habitat quality and nutrient concentrations were correlated with macroinvertebrate community metrics. They also found that forested streams, such as Lusk Creek, had the richest macroinvertebrate communities. Past work and need for further study. The Illinois Environmental Protection Agency (IEPA) and Illinois Department of Conservation (IDOC), now Illinois Department of Natural Resources (IDNR), have conducted limited investigations of Lusk Creek. The IEPA and IDNR are responsible for assessing stream quality and stream fisheries, respectively (Hite and Bertrand 1989). As part of a continued statewide effort, IEPA biologists periodically have sampled the benthic macroinvertebrate community in Lusk Creek to monitor stream quality (Hite et al. 1990). The Illinois Natural History Survey (INHS) is responsible for recording Illinois’ biological diversity (INHS 2008a). Since the 1930s, INHS scientists have sampled Lusk Creek sporadically, focusing on the Ephemeroptera (mayflies), Plecoptera (stoneflies), and Trichoptera (caddisflies) (EPT) fauna as part of a statewide inventories program (INHS 2008b). In doing so, they have documented 65 EPT species, making it the most EPT-rich stream in the state (Thomas 2001). Limited studies to date suggest Lusk Creek may support high stream insect diversity. However, most studies on the drainage have been system- atic in nature (e.g., Frison 1935, Ross 1944, Burks 1953, Poulton and Stewart 1991, Moulton and Stewart 1996) or have been conducted as part of statewide monitoring (e.g., Hite et al. 1990, Shasteen et al. 2003). Thus, a more thorough baseline study of the stream insects and habitat features of this system would be valuable for comparison with future surveys and for establishing regional reference conditions for Illinois streams. Identification of reference sites and conditions is important in the development of stream biological assessment programs and, thus, is a major focus of the USEPA and other agencies (USEPA 1995). Lusk Creek is important in this regard in that it provides the rare opportunity to examine physical and biological attributes of a relatively unimpaired drainage within a highly disturbed Illinois landscape. 2013 THE GREAT LAKES ENTOMOLOGIST 47

Study objectives. The objectives of this investigation were to: (1) survey the aquatic insects within the upper portion of the Lusk Creek watershed, (2) examine variation in the physical and chemical properties of the upper reaches, (3) use landscape features to explain patterns of insect distributions among sites, and (4) use observed taxa richness to estimate watershed-scale richness.

Materials and Methods This study was conducted from May 2003 to April 2005 in the upper portion of the Lusk Creek watershed. This watershed lies within the Shawnee Hills Natural Division (SH) of Illinois and is divided into the Greater SH and Lesser SH (Fig. 2). The Greater SH has sandstone geology with high topographic relief, and the Lesser SH has limestone and sandstone geology with lower topographic relief (Schwegman 1973). A digital map of the Lusk Creek watershed and its stream was used in conjunction with GIS ArcView 3.2 (ESRI 1999) to determine various features of the study area. The map included the NLCDS, National Elevation Data Digital Elevation Model with a 10-m resolution (NED) (Gesh et al. 2002), and USGS National Hydrography Dataset (NHD) (USGS 2003). Specifically, the NED was used to determine the watershed boundaries and watershed areas; the NLCDS and watershed boundaries were used to determine acreage by land cover type; the watershed areas and land acreage to determine watershed land cover composition; and the NHD was used to determine stream order, drainage density, and main channel sinuosity. Stream order was assigned according to Strahler (1963). Field measurements of basic physicochemical features of each site were made occasionally throughout the study period (Turner 2012). Study sites. In May 2003, potential study sites were identified using the United States Geological Survey National Land Cover Data Set (NLCDS) (Vogelmann et al. 2001), which delineated vegetation types found in the Lusk Creek watershed. Streams with watersheds that were predominately forested were selected for further investigation. These sites were located with a compass and 7.5” series quadrangle maps (USFS 1996). On-site evaluation criteria included the presence of riparian forest, stable banks, and embedded substrata; all criteria are considered representative of high quality stream habitat (Barbour et al. 1999). We also included the absence of filamentous algae and insects as dominant taxa in our evaluation. Originally, eight sites were selected. One was on Lusk Creek, itself (i.e., near Dog Hollow [L@DH]); six were on Lusk Creek tributaries (i.e., Bear Branch [BB], Copperous Branch [CB], Dog Hollow [DH], Little Bear Branch [LBB], Ramsey Branch [RB], and Little Lusk Creek, upstream of the confluence with East Fork [LL@EF]); and one was on East Fork [EF], a Little Lusk Creek tributary) (Fig. 3). All sites were first- and second-order streams. Three additional sites, higher-order segments, were added subsequently (i.e., Lusk Creek, downstream of the Lusk Creek Canyon Nature Preserve [L@LL] [third-order]; Little Lusk Creek, tributary of Lusk Creek, downstream of Martha’s Woods [LL@L] [third-order]; and Lusk Creek at the Eddyville Blacktop [L@Rd] [fourth-order], this latter site because earlier collections suggested high insect diversity (Fig. 3). The 11 study sites were located in the upper portion of the Lusk Creek stream system, nine in the Greater SH and two (CB, L@Rd) in the Lesser SH (Figs. 2 and 3). Physiochemical features. Watershed area of the sites ranged from 2.07–110.91 km2 and base flow ranged from no flow to 131.63 m /sec on sample dates (Appendix 1). Sites that had only isolated pools or were completely dry in September 2003 were considered intermittent; those with flow at this time were considered perennial. All first-order streams (LBB, RB, DH) were intermittent, 48 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Fig. 3. Lusk Creek watershed with sampling sites and their subbasins. 2013 THE GREAT LAKES ENTOMOLOGIST 49 and, with one exception (CB), all higher-order streams (2–4) were perennial (Appendix 1). Forest was the dominant watershed land cover type followed by grass and row crop (Appendix 2). Most sites had more than 75% forest and less than 20% grass and 7% row crop. RB was the only site with less than 50% forest and more than 20% row crop. Drainage density values generally were low, ranging from 1.20 to 2.04 km/ km2 (Appendix 3). Hillside slope values also generally were low, ranging from 1 to 15%; most sites were 3% or less (Appendix 3), with the three first-order streams (LBB, RB, DH) and the smallest second-order stream (BB) having the highest slopes. Stream sinuosity ranged from 1.17 to 3.36 in first- to third-order streams and 4.96 in the only fourth-order stream (L@Rd) (Appendix 3). All sites had substrata dominated by cobble and gravel, comprising 67–91% of all materials combined. All sites, with one exception (L@Rd), had bedrock/boulder as the third most common substratum; L@Rd had sand (Appendix 4). Canopy cover ranged from 9 to 72%, and eight sites had values ranging from 28 to 45% (Appendix 4). LBB, a first-order stream, had the highest value and was the only heavily shaded site (Appendix 4). Most sites had more erosional habitat than pool habitat, with the exception of LL@EF and RB, with 46 and 28% erosional habitat, respectively (Appendix 5). Water chemistry measurements were taken at 4–6-week intervals from June 2003 to April 2004. Warmest temperatures generally were recorded during August 2003; the coldest were during January/February 2004 (Appendix 6). Most pH measurements fell within the ideal range for aquatic insects (i.e., 6.5–8.0; USEPA 2011), ranging from 5.78 (DH, August 2003) to 8.12 (CB, February 2004) (Appendix 7). Although some of the conductivity measurements were within the ideal range for aquatic biota (i.e., 150–500 µS/cm; USEPA 2011), most were not (Appendix 8). Measurements ranged from 51.9–487.2 µS/cm, with 50 (81.0%) measurements below 150 µS/cm (Appendix 8). Invertebrate sampling. Benthic samples were collected during June 2003 with 10 sampling units per site. The sampling effort was proportional to the percentages of riffle and bank habitat present, which were estimated visually. If undercut bank habitat represented less than 10% of the habitat area at a site (10 of 11 sites), then only riffle habitats were sampled. For the eleventh site (LL@EF), riffles represented 90% and bank 10% of the total habitat, so riffle and bank habitats were sampled proportionally. All riffle sampling units were collected at approximately 10-meter increments. Riffle habitats were sampled using a Surber sampler (following Hauer and Resh 2007) with a 363-micron mesh and a 0.09-m2 frame. The sampler frame was placed on the substrata, and large, moveable rocks were washed in front of the sampler and then set outside the frame. The remaining substrata were disturbed thoroughly. The contents of the sampler were placed in a 34 × 25-cm white tray, rinsed with 95% ethanol to remove most of the sediments and placed in 95% ethanol. At each site, all riffle sample units were combined into one sample and represented a variety of substrate types and flow regimes. At the LL@EF site, the undercut bank habitat was sampled using a 0.3-m wide standard D-frame net with a 500-micron mesh. The net was placed under the bank and a 0.09-m2 area was disturbed thoroughly. Disturbed roots were cut and placed in the net. The net contents were placed in a 34 × 25-cm white tray and transferred to 95% ethanol. For this site, the bank sample was kept separate from the combined riffle sample. Each of the combined riffle samples for the 10 sites and the riffle and bank sample for the LL@EF site were labeled per site and kept separate. Within 24 hours and again in 2 weeks, the old ethanol was replaced with fresh 95% ethanol. 50 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Sample processing. For examination and analyses, two slightly different subsamples were collected. For the 10 sites consisting only of a combined riffle sample, the subsample comprised one half of the combined sample. For the LL@EF site, the subsample comprised 4/9ths of the combined sample plus the undercut bank sample. To obtain a subsample, the material was drained of alcohol using a sieve with a 250-micron mesh and rinsed in tap water. For sites with 10 combined riffle samples, the material was placed in a 37 × 21-cm white metal tray with a 5 × 2 grid drawn on the bottom and spread evenly over 10 sections. For the site with nine combined riffle samples, the material was spread over nine sections. The contents on each grid section were placed in a pint mason jar with 95% ethanol. Each grid section corresponded to a number, one through ten. To obtain the riffle subsample, numbers were selected randomly. If ten riffles were sampled, five numbers were selected; if nine, four were selected. For each subsample, a small amount of material was placed in a dish half- filled with 70% ethanol, and the animals were examined using a microscope with 10× magnification or, if needed, up to 45× magnification and removed. Animals were separated by order and placed in additional dishes filled with 70% ethanol. Most insect taxa subsequently were identified to the generic level (following Merritt and Cummins 1996) and functional feeding groups were assigned (following Merritt et al. 2008). However, chironomids (Diptera) were identified only to subfamily or tribe and dytiscids, hydrophilids (Coleoptera), and doli- chopodids (Diptera) only to family. Once identified, the insects were placed in vials with 70% ethanol. Diversity measurements and nonparametrics statistics. Biological diversity is the number of taxa as well as their relative abundance in the community (Lloyd and Ghelardi 1964). The number of taxa also is known as richness (McIntosh 1967). The relative abundance of each taxon describes the degree of community evenness (i.e., the more equal the value, the higher the evenness) (Smith and Smith 2001). Many investigators have used the term diversity to mean only richness. Therefore, it is important to understand how the term is being used (Hayek and Buzas 1997). For the present study, biological diversity refers to a mathematical derivation of both richness and evenness. All taxonomic units and all individuals were treated as equals, and abundance measurements were made consistently. Samples were assumed to represent continuous distributions. Nonparametric diversity indices and richness estimators were used to describe the aquatic insect community. The Shannon diversity index (H´) was used as a measure of diversity and calculated using the equation H´ = -∑pi ln pi, where p is the proportion for the ith taxa (Magurran 2004). H´ was calculated for each site. Variance also was calculated (following Hayek and Buzas 1997). The first-order jackknife was used to estimate taxa richness (observed and unobserved) from a single data set (following Manly 1997). This procedure was used because the number of taxa in a sample usually is an underestimate of the true taxa richness (Manly 1997). Palmer (1990, 1991) found the first-order jackknife to be the most precise and least biased of (Palmer 1990) or less biased than other (Palmer 1991) nonparametric diversity estimators.

Chao1 has been used to estimate taxa richness (Colwell and Coddington 1994). It was developed to estimate the total the number of classes (observed and unobserved) (Chao 1984). For this study, Chao1 was calculated for all sites 2 combined (following Colwell and Coddington 1994). Chao1 is equal to Sobs + ((F1 )/ (2F2)), where Sobs is the observed number of taxa, F1 represents the singletons (one individual ), and F2 represents the doubletons (two individuals). Variance also was calculated (following Chao 1987). 2013 THE GREAT LAKES ENTOMOLOGIST 51

An alternative form, Chao2, uses presence/absence information (Colwell and Coddington 1994). For this study, Chao2 was calculated for all sites combined 2 (following Colwell and Coddington 1994). Chao2 is equal to Sobs + ((Q1 )/(2Q2)), where Sobs is the observed number of taxa, Q1 represents the uniques (found at only one site) and F2 represents the duplicates (found at two sites). Variance also was calculated (following Chao 1987). Sampling at the LL@EF site included both riffle and undercut bank habitats. For all other sites, only riffles were sampled. Therefore, the LL@EF site was not comparable to the other sites and was not included in the analyses. Collection data from LL@EF are given in Turner (2012, Appendix A). Presented here are the results of the insect survey of Lusk Creek supple- mented with published life history information for selected taxa and preliminary analyses of taxa richness based on this survey.

Results and Discussion A total of 20,888 specimens, mostly immatures, were examined during the study and represented eight orders (Table 1). The Diptera, by far, was the most common order, with 18,590 specimens (89.0%), almost all of which were members of the Chironomidae and Simuliidae. The EPT (Ephemeroptera, Plecoptera, Trichoptera) combined were common with 1,550 specimens (7.4%). The Coleoptera was represented by 647 specimens, almost all of which were members of Stenelmis (Elmidae) (n = 612; 94.6%). The order Ephemeroptera was represented by at least 12 taxa in 11 genera and five families: Baetidae, Caenidae, Heptagenidae, Isonychiidae, and Leptophlebiidae (Table 1). Of the 11 genera, naiads of Acerpenna (Baetidae), Plauditus (Baetidae), and Habrophleboides (Leptophlebiidae) were the most numerous (Table 1). Acerpenna (formerly Baetis [McCafferty and Waltz 1990, McCafferty 1996]) is widespread in North America (Waltz and Burian 2008) and represented by three species (Purdue University 2011). The species occur in erosional lotic habitats and are both swimmers and clingers and collector-gatherers (Waltz and Burian 2008). Two species occur in Illinois [i.e., A. macdunnoughi (Ide), A. pygmaea (Hagen)] (Morihara and McCafferty 1979, Randolph and McCafferty 1998). Burks (1953), in his study of Illinois mayflies, stated that A. pygmaea prefers “small rivers or creeks with fairly rapid flow, such as Salt Fork River and Lusk Creek.” He reported A. pygmaea from a branch of Clear Creek (Union County) and the town of Herod (Pope County). A. macdunnoughi now also is known from Lusk Creek (INHS 2012b). A total of 242 naiads of Acerpenna (primarily A. macdunnoughi) was collected with most found at L@Rd (n = 190, 78.5%) and L@LL (n = 40, 16.5%) (Table 1), a fourth- and third-order perennial stream, respectively (Appendix 1). L@Rd and L@LL have riffle areas of 90 and 88%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the naiads among the sites (Table 1) showed a distinct preference for L@LL and L@Rd, third- and fourth-order sites, respectively. Plauditus (formerly Pseudocloeon, then Baetis [McCafferty and Waltz 1990, Lugo-Ortiz and McCafferty 1998]) is widespread in North America and represented by 10 species (Waltz and Burian 2008). The species occur in lotic habitats (erosional and depositional) and are both swimmers and clingers and collector-gatherers (Waltz and Burian 2008). Edmunds et al. (1976) reported that Pseudocloeon naiads are found in all stream sizes but predominately in shallow, fast water, often riffles. 52 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 g 1 4 2 4 6 13 25 24 242 295 Total c 2 3 1 31 L@ 190 Rd c 1 9 2 5 40 11 27 17 L@ LL c 6 1 1 1 89 14 LL@L c 4 1 1 28 LL@EF c a 2 1 1 2 72 L@DH c 8 EF Study Sites No. Specimens b 14 CB c 1 4 BB b 16 DH b 10 RB b LBB . . . . (Say) spp spp. spp spp. spp spp femoratum def Acerpenna Baetis flavistriga McDunnough

Taxon Ephemeroptera Baetidae B. intercalaris McDunnough Plauditus Procloeon Caenidae Caenis spp. Heptageniidae Leucrocuta Stenacron Stenonema Isonychiidae Isonychia Table 1. Number of specimens per taxon collected from riffle habitats study sites in the 2003 survey Lusk Creek Pope County, Illinois. 2013 THE GREAT LAKES ENTOMOLOGIST 53 g 2 1 2 15 33 142 117 227 Total c 2 4 6 L@ Rd c 7 1 75 12 11 L@ LL c 7 10 18 LL@L c 1 2 11 21 11 LL@EF c a 7 1 3 46 22 L@DH c 1 1 13 36 EF Study Sites No. Specimens b 2 3 4 3 CB c 1 2 1 1 52 29 BB b 1 1 1 10 36 DH b 1 1 RB b 5 2 54 LBB

. spp h . . spp spp

spp h def Amphinemura Amphinemura Perlesta

Perlidae Taxon Leptophlebiidae Habrophleboides spp. Paraleptophlebia spp. Odonata Coenagrionidae Plecoptera Capniidae Allocapnia Leuctridae Nemouridae Neoperla Table 1. Continued. 54 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 g 7 1 3 1 89 251 Total c 1 1 4 25 L@ Rd c 1 11 80 L@ LL c 3 39 LL@L c 22 15 LL@EF c a 2 1 16 45 L@DH c 1 6 13 EF Study Sites No. Specimens b 2 14 CB c 1 18 14 BB b 3 DH b 2 7 RB b 1 1 3 LBB

. . spp (L.) . spp spp . def cornutus Cheumatopsyche Corydalus Sialidae Taxon Hemiptera Veliidae Microvelia Megaloptera Corydalidae Nigronia Sialis Trichoptera Glossosomatidae Agapetus illini Ross Hydropsychidae spp Table 1. Continued. 2013 THE GREAT LAKES ENTOMOLOGIST 55 g 3 3 1 2 1 69 19 30 15 Total c 2 2 7 L@ Rd c 1 9 12 11 L@ LL c 3 1 1 3 2 39 LL@L c 1 LL@EF c a 5 4 11 L@DH c 4 4 1 EF Study Sites No. Specimens b 1 1 CB c 1 6 BB b 1 DH b 2 3 RB b 5 LBB

spp. spp. h spp.

Ross (Ross) shawnee Neotrichia Ross (Walker) def Wormaldia Rhyacophila R. glaberrima

Polycentropus Polycentropus Leptoceridae Rhyacophilidae Taxon Hydroptilidae Hydroptila Philopotamidae Chimarra feria C. obscura Polycentropodidae fenestra Ulmer Table 1. Continued. 56 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 g 1 1 1 5 5 13 15 612 101 Total c 1 1 6 L@ 120 Rd c 4 14 L@ 185 LL c 1 1 3 63 LL@L c 4 60 30 LL@EF c a 1 1 5 33 L@DH c 1 9 32 EF Study Sites No. Specimens b 8 1 23 15 CB c 1 1 7 70 BB b 2 2 11 DH b 4 4 4 11 RB b 3 4 1 1 6 LBB

spp

spp. spp. spp. (DeKay)

h complex complex herricki def McLachlan concinnus Helichus Ectopria thoracica thoracica Ectopria Psephenus Neophylax

Dytiscidae Ceratopogonidae

Taxon Uenoidae

Coleoptera

Dryopidae

Elmidae Stenelmis Hydrophilidae Psephenidae

(Ziegler)

Diptera

Atrichopogon Bezzia Table 1. Continued. 2013 THE GREAT LAKES ENTOMOLOGIST 57 g 1 1 3 1 11 45 29 420 Total 8473 3072 1601 c 1 4 26 L@ 735 190 226 Rd c 7 38 L@ 350 385 LL 1136 c 4 17 628 337 132 LL@L c 1 1 1 12 76 658 377 LL@EF c a 2 160 245 1693 1281 L@DH c 1 19 60 199 177 EF Study Sites No. Specimens b 4 52 11 505 146 173 CB c 1 29 70 61 BB 1039 b 1 30 33 40 747 103 DH b 5 2 3 1 17 26 891 197 RB b 1 20 15 13 242 LBB

h h h

h h spp. sp. sp. spp h def Anopheles Hemerodromia Psychoda Tanytarsini

Chironomini Culicidae Chelifera Psychodidae Taxon Dasyhelea Chironomidae Tanypodinae Orthocladiinae Chironominae

Dixidae Dolichopodidae Empididae

spp.

Table 1. Continued. 58 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 g 5 1 2 92 19 22 60 Total 4686 20,888 c 1 27 L@ Rd 1837 3429 c 3 1 1 1 33 L@ 666 LL 3134 c 2 8 1 30 570 2005 LL@L c 5 3 16 26 141 1503 LL@EF c a - 3 1 9 30 577 4252 L@DH c 3 2 29 23 273 893 EF Study Sites No. Specimens b 2 2 2 23 216 CB 1204 c 6 1 3 26 355 BB 1775 b 9 1 25 20 DH 1073 b 6 2 32 22 RB 1231 b 1 1 10 20 389 LBB . . sp . spp. spp. spp spp. spp. spp

i def Hexatoma Limonia Molophilus Pilaria Tipula

Simuliidae Dicranota

Taxon Simulium Tipulidae

Total no. specimens Total no. specimens Total taxa Unidentified genus or genera. Specimens collected during June 2003. See Appendix 1 for complete names and stream order. Intermittent stream. Total number of specimens per taxon. All specimens were immatures (i.e., naiads, nymphs, larvae) except for Hemiptera and Perennial stream. For purpose of this table, spp. treated as one taxon. Minimum number of taxa per site and for all sites combined. Table 1. Continued. a b c d e some Coleoptera as follows: Veliidae, adults and nymphs; Dryopidae, adults; Dytisci dae, adult; Elmidae, adults and larvae. f g h i 2013 THE GREAT LAKES ENTOMOLOGIST 59 Three species occur in Illinois [i.e., P. armillatus (McCafferty and Waltz), P. dubius (Walsh), P. punctiventris (McDunnough)] and two in southern Illinois (i.e., P. dubius, P. punctiventris) (Burks 1953, Randolph and McCafferty 1998). Burks (1953) stated that P. dubuis and P. punctiventris prefer “small rivers or creeks with fairly rapid flow, such as Salt Fork River and Lusk Creek.” He reported P. dubuis from Hutchins Creek and Lusk Creek and P. punctiventris from Hutchins Creek (Union County). A total of 295 naiads of Plauditus was collected from all sites except LBB and BB with most found at LL@L (n = 89, 30.2%) and L@DH (n = 72, 24.4%) (Table 1), a third- and second-order perennial stream, respectively (Appendix 1). LL@L and L@DH have riffle areas of 79 and 89%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the naiads among the sites (Table 1) showed a broad tolerance of stream order ranging from first-order to fourth-order sites. Habrophleboides is found in the eastern and midwestern United States and represented by four species (Waltz and Burian 2008). The species occur in lotic habitats (erosional and depositional); are swimmers, clingers, and sprawlers; and scrapers and collector-gatherers (Waltz and Burian 2008). They occur in streams with slow to moderately fast current and may be found in riffles although they occur more typically with submerged plants and woody debris (Edmunds et al. 1976). One species, H. americana (Banks), occurs in Illinois (Burks 1953, Ran- dolph and McCafferty 1998). Burks (1953) reported that H. americana prefers “small, temporary pools, usually along stream margins, which have greatly reduced or no current,” and reported it from the town of Herod (Pope County). A total of 142 naiads of Habrophleboides was collected from all sites except LBB, RB, and L@Rd (Table 1) with most found at BB (n = 52, 36.6%) and L@DH (n = 46, 32.4%) (Table 1), both second-order perennial streams (Appendix 1). BB and L@DH have riffle areas of 59 and 89%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the naiads among the sites, including a few specimens from two intermittent streams (DH and CB) (Table 1), showed a broad tolerance of stream order but differed from Acerpenna and Plauditus in that no specimens were collected at L@Rd. The order Plecoptera was represented by at least five taxa in four genera and four families: Capniidae, Leuctridae, Nemouridae, and Perlidae (Table 1). Of the four genera, naiads of Allocapnia (Capniidae) and Perlesta (Perlidae) were the most numerous (Table 1). Allocapnia is found in eastern North America (Stewart and Stark 2008) and represented by 47 species (DeWalt et al. 2012). The species are clingers and shredder-detritivores (Stewart and Stark 2008). Naiads of some species avoid summer temperatures by burrowing and entering diapause. They can be found at a depth of 10–20 cm and difficult to find (Harper and Hynes 1970). Eight species occur in Illinois [i.e., A. forbesi Frison, A. granulata (Claas- sen), A. mytica Frison, A. nivicola (Fitch), A. recta (Claassen), A. rickeri Frison, A. smithi Ross and Ricker, A. vivipara (Claassen)] (DeWalt et al. 2005), five of which are found in the Shawnee Hills (i.e., A. vivipara, A. rickeri, A. mytica, A. forbesi, and A. smithi) (Webb 2002, DeWalt et al. 2005). A. vivipara is the most common Illinois winter stonefly, whereas A. rickeri is one of the most common in the Shawnee Hills. A. mytica, A. forbesi, and A. smithi are restricted to the Shawnee Hills with the latter two found only on the eastern side (Webb 2002). A. vivipara is found in a variety of stream sizes and conditions, including organic enrichment, whereas the other four species are found in clear, cool, typically spring-fed streams with course substrata (Ross and Ricker 1971). A. forbesi can be found in streams that may experience summer drying (Ross and Ricker 1971). A. vivipara is known to undergo naiadal diapause (Harper and Hynes 1970). 60 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

A total of 117 naiads of Allocapnia was collected in June from all sites except LL@L. They were small and in diapause with the head bent over the body as described by Harper and Hynes (1970). They were collected at all sites except LL@L with most found at L@LL (n = 75, 64.1%) and LL@EF (n = 21, 17.9%) (Table 1), a third- and second-order perennial stream, respectively (Appendix 1). L@LL and LL@EF have riffle areas of 88 and 46%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the naiads among the sites, including occurrence in both intermittent and perennial streams (Table 1), showed a broad tolerance of stream order but a distinct preference for L@LL, a third-order site. Perlesta is widespread in North America (Stewart and Stark 2008) and represented by 30 species (DeWalt et al. 2012). Most species occur in lotic habitats (erosional and depositional), are clingers, and are engulfing predators and facultative collector-gatherers (primarily in early instars) (Stewart and Stark 2008). Nine species occur in Illinois [i.e., P. cinctipes (Banks), P. decipiens (Walsh), P. golconda DeWalt & Stark, P. lagoi Stark, P. ouabache Grubbs and DeWalt, P. shawnee Grubbs & Stark, P. shubuta Stark, P. teaysia Kurchner and Kondra- tieff, P. xube Stark & Rhodes)] (DeWalt and Grubbs 2011, DeWalt et al. 2012), four of which have been reported from Pope County (i.e., P. golconda, P. lagoi, P. shawnee, P. xube) (DeWalt et al. 2001, Grubbs 2005). P. golconda typically is found in large rivers (DeWalt et al. 2001, 2005). P. lagoi is the second most common species in Illinois and found throughout the state in small streams (DeWalt et al. 2001). P. shawnee is restricted to the southern unglaciated region of the state (DeWalt et al. 2001, 2005). P. xube is uncommon and found in small forested streams that may be reduced to pools in the summer (DeWalt et al. 2001). None of the species can be identified as naiads. A total of 227 naiads of Perlesta was collected from all sites with most found at LBB (n = 54, 23.8%), DH (n = 36, 15.9%), and EF (n = 36, 15.9%) (Table 1), the first two sites, first-order intermittent streams, the third site, a second-order perennial stream (Appendix 1). LBB, DH, and EF have riffle areas of 58, 69, and 85%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the naiads among the sites (Table 1) showed a broad tolerance of stream order but no obvious preference for any one site. The order Megaloptera was represented by at least three taxa in three genera and two families: Corydalidae and Sialidae (Table 1). Of the three genera, larvae of Nigronia (Corydalidae) were the most numerous (Table 1). Nigronia is found in eastern and central North America and represented by two species (Flint et al. 2008), N. fasciatus (Walker) and N. serricornis (Say) (Tarter et al. 1976). The species occur in lotic habitats (erosional and depositional), are clingers, climbers, and burrowers and are engulfing predators (Flint et al. 2008). Nigronia fasciatus and N. serricornis occur in Illinois (Tarter et al. 1976). Neunzig (1966) reported that N. fasciatus is restricted to “small, cool woodland streams,” whereas N. serricornis inhabits “large woodland streams” and “certain portions of rivers.” Tarter et al. (2006) found N. fasciatus most often in first-order streams and N. serricornis in second- to fourth-order streams but sometimes found the two species together. They reported the names of those streams but not the stream order. A total of 89 larvae of Nigronia (including fiveN. serricornis) was collected from all sites except LBB, RB, and DH (Table 1). Therefore, they were found only in second- to fourth-order streams (Appendix 1) with most found at LL@ EF (n = 22, 24.7%) (Table 1). LL@EF has a riffle area of 46% (Appendix 5) and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance for stream order, excluding first-order, but no obvious preference for any one site. 2013 THE GREAT LAKES ENTOMOLOGIST 61 The order Trichoptera was represented by at least 12 taxa in 11 genera and eight families: Glossosomatidae, Hydropsychidae, Hydroptilidae, Leptoceridae, Philopotamidae, Polycentropodidae, Rhyacophilidae, and Uenoidae (Table 1). Of the 11 genera, larvae of Cheumatopsyche (Hydropsychidae) and Chimarra (Philopotamidae) were the most numerous (Table 1). Cheumatopsyche is widespread in North America and represented by 44 species (Morse and Holzenthal 2008). The species occur in lotic erosional habitats, particularly in warmer streams and rivers, are clingers (net-spinners) that build fixed retreats, and are collector-filterers (Morse and Holzenthal 2008). They are found in moderate currents and build nets with intermediate mesh sizes (Wiggins 1996, 2004). In the Interior Highlands, streams may contain several congeners (Moulton and Stewart 1996). Nine species occur in Illinois [i.e., C. analis, (Banks), C. aphanta Ross, C. burksi Ross, C. campyla Ross, C. lasia Ross, C. oxa Ross, C. pasella Ross, C. sordida (Hagen), C. speciosa (Banks)] (Ross 1944), including three that occur in the Illinois Ozarks of the Interior Highlands (i.e., C. analis, C. campyla, C. oxa) (Moulton and Stewart 1996). C. campyla is found in a variety of environmental conditions (Moulton and Stewart 1996) including those not tolerated by other caddiflies (Ross 1944). It prefers larger streams (Ross 1944). C. analis is the most common species in the Interior Highlands (Moulton and Stewart 1996) and also tolerates degraded water quality (Ross 1944). C. oxa is found in small to medium streams (Moulton and Stewart 1996), particularly in small, spring- fed streams (Ross 1944). A total of 251 larvae of Cheumatopsyche was collected from all sites with most found at L@LL (n = 80, 31.9%) (Table 1), a third-order perennial stream (Appendix 1). L@LL has a riffle area of 88% (Appendix 5) and abundant cobble/ gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order but a distinct preference for L@LL, a third-order site. Chimarra is widespread in North America and represented by 21 species (Morse and Holzenthal 2008). The species occur in erosional lotic habitats in warm rivers and are clingers (saclike, silk net makers) and obligate collector- filterers (Morse and Holzenthal 2008). As with other philopotamids,Chimarra spp. are restricted to flowing waters (Wiggins 2004) that serve to inflate their silken filtering nets (Wiggins 1996). They use their membraneous labrum to remove fine particles from the inside of the nets (Wiggins 2004). Four species occur in Illinois [i.e., C. aterrima Hagen, C. feria Ross, C. obscura (Walker), C. socia Hagen] (Ross 1944), including two in the Illinois Ozarks in the Interior Highlands (i.e., C. feria , C. obscura) (Moulton and Stewart 1996). C. feria and C. obscura are common species in the Interior Highlands and often collected together (Moulton and Stewart 1996). C. feria is common in clear, fast streams in southern Illinois where it tolerates summer drying by seeking refuge in damp conditions under rocks (Ross 1944). C. obscura also prefers clear, fast streams (Ross 1944). Both C. feria and C. obscura were collected in the present study, C. feria being the most common (Table 1). For C. feria, 69 larvae were collected with most found at LL@L (n = 39, 56.5%) (Table 1); for C. obscura (Walker), 19 larvae were collected with most found at L@LL (n = 11, 57.9%) (Table 1). LL@L and L@LL are third-order perennial streams (Appendix 1). LL@L and L@LL have riffle areas of 79 and 88%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the larvae of C. feria among the sites (Table 1) indicates a broad tolerance of stream order, although the larvae of both C. feria and C. obscura showed a distinct preference for the downstream reach of the third-order sites (see above). 62 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

The order Coleoptera was represented by at least six taxa in four genera and five families: Dryopidae, Dytiscidae, Elmidae, Hydrophilidae, and Psephenidae (Table 1). Of the four genera, larvae of Stenelmis (Elmidae) were the most numerous (Table 1). Stenelmis is widespread in North America and represented by 33 species (White and Roughley 2008). The species occur in erosional lotic habitats with coarse sediments and detritus and are clingers and both scrapers and collector- gatherers (White and Roughley 2008). Several species may be found together (Sanderson 1953). Three species are found in Illinois [i.e., S. crenata (Say), S. decorata Sand- erson, and S. vittipennis Zimmermann] (Brown 1983). In Wisconsin, S. crenata adults inhabit a variety of stream types, and S. decorata adults inhabit medium to large streams (Hilsenhoff and Schmude 1992). In Indiana, S. vittipennis is found in a variety of stream sizes (McMurray and Newhouse 2006). S. crenata has been reported from Lusk Creek (Shasteen et al. 2003). A total of 612 larvae and adults of Stenelmis was collected from all sites with most found at L@LL (n = 185, 30.2%) and L@Rd (n = 120, 19.6% (Table 1), a third-order and fourth-order perennial stream, respectively (Appendix 1). L@LL and L@Rd have riffle areas of 88 and 90%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The general increase in numbers moving downstream (Table 1) showed a distinct preference for L@LL and L@Rd, third- and fourth-order sites, respectively. The order Diptera was represented by at least 21 taxa in 15 genera and nine families: Ceratopogonidae, Chironomidae, Culicidae, Dixidae, Dolicho- podidae, Empididae, Psychodidae, Simuliidae, and Tipulidae (Table 1). Of the nine families, larvae in the Ceratopogonidae, Chironomidae, Simuliidae, and Tipulidae were the most numerous (Table 1). Ceratopogonidae. In the present study, this family was represented by three genera. Of these, the larvae of the Bezzia complex were the most numerous. The Bezzia complex includes at least Bezzia and Palpomyia. There is difficulty in separating these two genera (Courtney and Merritt 2008), but the species appear to occur in different habitats (Merritt and Webb 2008). Thus for this discussion, they are treated separately. Bezzia is widespread in North America and represented by 52 species (Merritt and Webb 2008). The species are found in lentic (littoral, profundal, and sometimes limnetic) and lotic (in hot springs [algal mats]) habitats. They are burrowers, occasionally planktonic (swimmers), and engulfing predators (Merritt and Webb 2008). Palpomyia is widespread in North America and represented by 31 species (Merritt and Webb 2008). The species are found in lotic (erosional and depositional [detritus]) and lentic (littoral, profundal, sometimes limnetic) habitats. They are burrowers, occasionally planktonic (swimmers), and predators (engulfers) and collector-gatherers (Merritt and Webb 2008). A total of 101 larvae of the Bezzia complex was collected at all sites combined with most found at LL@EF (n = 30, 29.7%) (Table 1), a second-order perennial stream (Appendix 1). Their preference for lotic habitats more closely resembles that of Palpomyia than of Bezzia. LL@EF has a riffle area of 46% (Appendix 5) and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order but no obvious preference for any one site. Chironomidae. This family was represented by four taxa (i.e., Tanypo- dinae, Orthocladinae, Chironomini, and Tanytarsini), the larvae of which were numerous (Table 1). 2013 THE GREAT LAKES ENTOMOLOGIST 63 Species of Tanypodinae are widespread in North America and represented by 39 genera (Ferrington et al. 2008). They occur in all lentic and lotic habitats and generally are sprawlers and swimmers and are predators (engulfers and piercers) (Ferrington et al. 2008). A total of 420 larvae of Tanypodinae was collected at all sites with the most found at L@DH (n = 160, 38.1%) (Table 1), a second-order perennial stream (Appendix 1). L@DH has a riffle area of 89% (Appendix 5) and abundant cobble/ gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order with a distinct preference for the L@DH site. Species of Orthocladiinae are widespread in North America, especially in the North, and represented by 82 genera (Ferrington et al. 2008). The subfamily is diverse and, consequently, species are found in a variety of habitats (Epler 2001). Larvae occur primarily in lotic habitats, but many occur in lentic habitats (primarily oligotrophic lakes) and generally are burrowers (tube-builders) and collector-gatherers and scrapers (Ferrington et al. 2008). They dominate in streams with coarse substrates and colder waters, typically low-order, headwater streams (Pinder 1995). A total of 8,473 larvae of Orthocladiinae was collected at all sites with most found at L@DH (n = 1,693, 20.0%) (Table 1), a second-order perennial stream (Appendix 1). L@DH has a riffle area of 89% (Appendix 5) and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order but a distinct preference for the BB, L@DH, and L@LL sites, all second- and third- order perennial streams. Species of Chironomini are widespread in North America and represented by 50 genera (Ferrington et al. 2008). They generally occur in lentic (littoral and profundal) and lotic (depositional) habitats and usually are burrowers and collector-gatherers (Ferrington et al. 2008). A total of 3,072 larvae of Chironomini was collected at all sites with most found at L@DH (n = 1,281, 41.7%) (Table 1), a second-order perennial stream (Appendix 1). L@DH has a riffle area of 89% (Appendix 5) and abundant cobble/ gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order but a distinct preference for the L@DH site. Species of Tanytarsini are widespread in North America and represented by 18 genera (Ferrington et al. 2008). They generally occur in lotic (erosional and depositional) and lentic (littoral) habitats and usually are burrowers or clingers (tube-builders) and collectors (gatherers and filterers) (Ferrington et al. 2008). A total of 1,601 larvae of Tanytarsini was collected at all sites with most found at L@LL (n = 385, 24.1%) (Table 1), a third-order perennial stream (Ap- pendix 1). L@LL has a riffle area of 88% (Appendix 5) and abundant cobble/ gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order. Simuliidae. This family was represented by only Simulium in the present study, the larvae of which were numerous (Table 1). Simulium is widespread in North America and represented by 154 species (Alder and Currie 2008). The species occur in lotic and lentic erosional habitats and are clingers and collector-filterers (Alder and Currie 2008). Although simuliids usually are found in moderate-sized streams, they can be found in smaller or larger streams (Crosskey 1990). A total of 4,686 larvae of Simulium was collected at all sites with most found at L@Rd (n = 1,837, 39.2%) (Table 1), a fourth-order perennial stream (Appendix 1). L@Rd has a riffle area of 90% (Appendix 5) and abundant cobble/gravel substrata 64 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

(Appendix 4). The general increase in numbers moving downstream (Table 1) showed a general preference for third- and fourth-order sites. Tipulidae. This family was represented by at least six species in six genera in the present study. Of the six genera, the larvae of Dicranota were the most numerous (Table 1). Dicranota is widespread in North America and represented by 55 species (Byers and Gelhaus 2008). The species are found in lotic (erosional and depositional [detritus]) and lentic (littoral [detritus]) habitats and along margins of both habitats. They are sprawlers and burrowers and engulfing predators (Byers and Gelhaus 2008). A total of 92 larvae of Dicranota was collected at all sites except LBB and L@Rd with most found at EF (n = 29, 31.5%) and DH (n = 25, 27.2%) (Table 1), a second-order perennial and first-order intermittent stream, respectively (Ap- pendix 1). EF and DH have riffle areas of 85% and 69%, respectively (Appendix 5), and abundant cobble/gravel substrata (Appendix 4). The distributional pattern of the larvae among the sites (Table 1) showed a broad tolerance of stream order with a distinct preference for the DH, EF, and LL@EF sites, the first, a first-order intermittent site, and the latter two, second-order perennial sites. Insect distribution patterns. The number of taxa per site ranged from 20 to 33 (Table 2). Comparing sites overall, there was a relationship between stream order, hydrologic status, and taxa richness. Generally, as stream order increased, streams transitioned from intermittent to perennial and taxa richness increased. Perennial streams generally had more EPT taxa than intermittent streams, a pattern not evident in Diptera, the only other well-represented group (Table 2). When stream conditions remain stable, habitat becomes more stable and stream biota more diverse (Hynes 1970). Feminella (1996) found that richness, both overall and the EPT, was related to hydrologic status, and more permanent streams had increased numbers of taxa. In this investigation, higher-order perennial streams had increased richness. Compared to intermittent streams, perennial streams would provide more stable hydrologic conditions that would allow for greater diversity. In the present study, species of the Chironomidae comprised three of the four most commonly collected taxa, the fourth being Simulium; thereafter, there was a sharp decrease in specimens collected (Table 3). Of those taxa identified to genus, the Tipulidae was the most diverse, with six genera (Table 1). Un- doubtedly, the Chironomidae would have far exceeded six had the specimens been identified to genus. Based on the number of specimens collected for all taxa (Table 3), the Orthocladiinae (n = 8,473; 40.6%), Chironomini (n = 3,072; 14.7%), and Tanytarsini (n = 1,601; 7.7%) were among the most commonly collected taxa, representing almost 63% of all taxa collected. Further, the number of specimens within the Orthocladiinae far exceeded the numbers within the other subfamilies, supporting Tokeshi’s (1995) statement that the Orthocladiinae is the most abundant subfamily of the chironomids in temperate streams of the Northern Hemisphere. The River Continuum Concept (RCC) prediction of higher diversity in medium-sized streams compared to headwater streams (Vannote et al. 1980) was moderately supported by this investigation. Most medium-sized reaches had higher diversity than the smaller headwater reaches (Table 2). In general, patterns of functional feeding structure supported RCC pre- dictions for first- to fourth-order streams (Vannote et al. 1980). Of the 60 taxa collected (Table 1), 32 were represented by one feeding group and included ten collectors (filterers, gatherers), 15 predators (engulfers, piercers), four scrapers (grazers), and three shredders; the remaining taxa were combinations of these categories (Table 4). Functional feeding groups were distributed among all stream orders (Table 5). 2013 THE GREAT LAKES ENTOMOLOGIST 65

Table 2. Stream order and number of taxa collected from riffle habitats of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois. Taxaa Study Stream Hydrologic Siteb Order Statusc No.Taxad EPT D C M H O

LBBe 1 I 20 5 9 4 1 1 0

DHe 1 I 20 8 10 2 0 0 0 RBe 1 I 22 6 13 2 0 1 0 CBe 2 I 23 8 12 2 1 0 0

EFe 2 P 23 9 10 2 1 1 0 LL@EFf 2 P 26 11 12 2 1 0 0 BBe 2 P 26 11 10 3 1 1 0 L@Rde 4 P 27 12 9 3 2 1 0 L@DHe 2 P 30 16 10 2 2 0 0 LL@Le 3 P 30 16 10 3 1 0 0 L@LLe 3 P 33 17 11 2 1 1 1 Total 60 29 20 6 3 1 1 aEPT = Ephemeroptera, Plecoptera, Trichoptera; D = Diptera; C = Coleoptera; M = Megaloptera; H = Hemiptera; O = Odonata. bSee Appendix 1 for complete names. cI = intermittent, P = perennial. dMinimum number of taxa per site and for all sites combined (see Table 1). eRiffle comprised entire sample. fRiffle and undercut bank sampled. 66 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 3. Abundant (>10%), common (1% to ≤10%), and rare (<1%) taxa collected from riffle habitats of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Taxon Total No. Specimens Overall Proportion (%)

Orthocladiinae 8,473 40.6 Simulium spp. 4,686 22.4 Chironomini 3,072 14.7 Tanytarsini 1,601 7.7 Stenelmis spp. 612 2.9 Tanypodinae 420 2.0 Plauditus spp. 295 1.4 Cheumatopsyche spp. 251 1.2 Acerpenna spp. 242 1.2 Perlesta spp. 227 1.1 Habrophleboides spp. 142 0.7 Allocapnia spp. 117 0.6 Bezzia complex 101 0.5 Dicranota spp. 92 0.4 Nigronia spp. 89 0.4 Chimarra spp. (n = 2) 88 0.4 Other rare taxa (n = 43) 380 1.8 Total 20,888 100.0 2013 THE GREAT LAKES ENTOMOLOGIST 67 2–4 3 3–4 1–4 3–4 2–3 2–3 2–3 2–4 3 1–3 1–2 3 1–4 1–2 2 1–4 2–4 1–4 Stream Order b Cg Cg/Sc(facultative) Cg/Sc(facultative) Cg Cg/Sc Cg/Sc Sc/Cg Sc/Cg(facultative) Sc/Cg(facultative) Cf/Pe Sc/Cg Cg/Sh(facultative) Pe Sh Sh Sh/Cg(facultative) Pe/Cg(facultative) Pe Pp Functional Feeding Group

spp. spp. sp. . femoratum spp.

spp. spp. spp. spp spp. spp. spp. spp. spp. spp. Tribe, Genus, or Species Acerpenna Baetis flavistriga Baetis intercalaris Plauditus Procloeon Caenis Leucrocuta Stenacron Stenonema Isonychia Habrophleboides Paraleptophlebia Allocapnia Amphinemura Perlesta Neoperla Microvelia Family or Subfamily Baetidae Caenidae Heptageniidae Isonychiidae Leptophlebiidae Coenagrionidae Capniidae Leuctridae Nemouridae Perlidae Veliidae Order Ephemeroptera Odonata Plecoptera Hemiptera Table 4. Functional feeding group(s) and associated stream orders for taxa collected from riffle habitats of sites used in 2003 insect survey Lusk Creek in Pope County, Illinois. 68 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 4 2–4 1–2 2 1–4 2, 4 3 3 2–4 3–4 2–3 1–3 1 1–3 2 2 1–4 1–4 1–3 Stream Order b

c Pe Pe Pe/Cg(one species) Sc/Cg(facultative) Cf Pc/Sc(facultative) Sc Cg/Cf/Sh/Sc/Pe Cf(obligate) Cf(obligate) Cf(obligate) Pe/Cf/Sh Pe/Sc/Cg/Sh Pe/Sc/Cg/Sh Sc(obligate) Pe Sc/Sh Sc/Cg Pe Functional Feeding Group

spp. spp. spp. spp. spp.

spp. spp. spp. Tribe, Genus, or Species Corydalus cornutus Nigronia Sialis Agapetus illini Cheumatopysche Hydroptila Neotrichia Chimarra feria Chimarra obscura Wormaldia shawnee Polycentropus Rhyacophila fenestra Rhyacophila glaberrima Neophylax concinnus Helichus Stenelmis Family or Subfamily Corydalidae Sialidae Glossosomatidae Hydropsychidae Hydroptilidae Leptoceridae Philopotamidae Polycentropodidae Rhyacophilidae Uenoidae Dytiscidae Dryopidae Elmidae Hydrophilidae Order Megaloptera Trichoptera Coleoptera Table 4. Continued. 2013 THE GREAT LAKES ENTOMOLOGIST 69 1 3–4 1–2 1–4 1–2 1–4 1–4 1–4 1–4 2 1 1 1–2, 4 2–4 1 1–4 1–3 1–4 Stream Order b

Pp Pp Sc Sc Cg/?Sc Pe Cg/Sc Pe/Pp Cg/Sc Cg Cf/Cg Cf Cg/?Cf Pp Cg Cf Pe Pe Functional Feeding Group

spp. spp. herricki spp.

spp. spp. sp. spp. spp. spp. complex Tribe, Genus, or Species Ectopria thoracica Psephenus Atrichopogon Bezzia Dasyhelea Chironomini Tanytarsini Anopheles Chelifera Hemerodromia Psychoda Simulium Dicronota Hexatoma

Family or Subfamily Psephenidae Ceratopogonidae Chironomidae Tanypodinae Orthocladiinae Chironominae Culicidae Dixidae Dolichopodidae Empididae Psychodidae Simuliidae Tipulidae

Order Coleoptera Diptera

Table 4. Continued. 70 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 1–2 3 1, 3 1–3 Stream Order = plant piercer; Pc b

= collector-gatherer; Cg

Sh ---- ?Pe Functional Feeding Group Sh(obligate)/Cg/ Sc(facultative)

a = collector-filterer; Cf = shredder. Sh sp. spp. spp. spp. Tipula Tribe, Genus, or Species Limonia Molophilus Pilaria = scraper; and Sc (2008), generally at generic level:

Tipulidae = predator-piercer; Family or Subfamily Pp = predator-engulfer; Order Diptera For purpose of this table, each spp. treated as one taxon. As defined and listed by Merritt et al. Merritt et al. (2008) listed functional feeding group for adults and larvae; only larvae collected in present study. Table 4. Continued. a b Pe c 2013 THE GREAT LAKES ENTOMOLOGIST 71 h 1 0 0 1 0 3–4 g 1 1 3 0 0 2–4 f 1 0 0 0 0 2–3 e 2 2 4 0 1 1–4 d 0 0 2 0 0 1–3 c 0 0 1 0 0 1, 3 b Stream Order 0 0 1 0 0 1–2,4 a 0 0 0 0 2 1 –2 0 0 1 0 0 4 0 0 1 1 0 3 by stream order for taxa collected from riffle habitats of sites used in 2003 insect survey

1 0 1 1 0 2 0 1 1 1 0 1 i Functional Feeding Group Collector-filterers Collector-gatherers Predators Scrapers Shredders Third- to fourth-order streams. First- to third-order streams. First- to second-order streams. First- to second- and fourth-order streams. Second- to fourth-order streams. First- to fourth-order streams. First- and third-order streams. Second- to third-order streams. Excludes combined feeding groups (see Table 4). Table 5. Number of functional feeding groups Lusk Creek in Pope County, Illinois. a b c d e f g h i 72 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Biological diversity (richness and evenness). The combined insect richness for the 11 sites was 60 taxa representing eight orders (Table 2). The combined EPT richness was 29 taxa (12 Ephemeroptera, 5 Plecoptera, and 12 Trichoptera [Table 1]). The richness of each order comprising the EPT was actually less than that of the Diptera with 20 taxa. The remaining four orders (i.e., Coleoptera, Megaloptera, Hemiptera, and Odonata) were represented by six taxa or less and had a combined richness of 11 (Tables 1 and 2). Species richness will be underestimated if specimens are identified only to morphospecies because, at this level, genera may contain more than one species (Rosenberg et al. 2008). For this investigation, only 13 taxa were identified to the species level, 36 to the generic level, and the remaining 11 to tribe, subfamily, or family level (Table 1). Unique taxa may be undetected at the generic level or higher. Therefore, the observed taxa richness may have been underestimated. The benthic insect taxa richness can be used to infer habitat quality (Bar- bour et al. 1999). The EPT richness is used to evaluate stream health (Lenat 1988). Heatherly et al. (2007) found that Lusk Creek and other forested streams in southern Illinois with low nutrient levels had higher insect taxa richness than non-forested Illinois streams with high nutrient levels. For impaired streams in the Kaskaskia River system, Stone et al. (2005) found at least 13 insect taxa and only four EPT taxa (three tolerant ephemeropterans, no plecopterans, and one trichopteran represented by two specimens). When compared to Stone et al. (2005), the results of this study indicate that Lusk Creek is a high quality stream with a rich benthic insect community. Delucchi (1988) categorized taxa as abundant (> 10%), common (1% - ≤ 10%), or rare (< 1%) based on total number of specimens collected. Using these same categories for the 60 taxa in the present study, there were three abundant, seven common, and 50 rare taxa (Tables 1 and 3). Biological communities typically are comprised of a few abundant species, some common species, and a majority of rare species (Magurran 2004). The observed results in the present study follow that general pattern and reflect low evenness. Longino et al. (2002) used different collection methods in various habitats to assess the richness of a tropical ant community. When they compared collecting techniques, they found that typically a single method yielded fewer taxa and high numbers of rare ones, some which might be common elsewhere. In the present study, only one habitat in one season was sampled using one method. Of the 50 rare taxa, some may have been common in other habitats. For example, in the present study, one specimen of Anopheles was collected in the riffle sample at the BB site, but several were collected in the supplemental bank samples (unpublished data). Also, one specimen of Neophylax was collected in the riffle sample at the L@DH site, but several were observed in the spring. The Shannon diversity index gave H´ values for individual sites that ranged from 1.07 to 2.01 (Table 6). Sites with the lowest (RB, 1.07) and the highest (EF, 2.01) values had low richness (Tables 1 and 2). Therefore, the H´ value difference was due to a change in evenness. This can be observed in the relative abundances of the four most common taxa (Tables 3 and 7). RB had one dominant and one moderately dominant taxon, whereas EF had three moderately dominant taxa; therefore, EF had a higher evenness (Table 7). Increases in richness and evenness will cause H´ values to increase (Lloyd and Ghelardi 1964). Therefore, it appears that differences among sites may be related to changes in richness, evenness, or both. The proportion of each taxon determines its influence on the index value. The dominant taxa con- tribute less than moderately dominant taxa, and the maximum contribution occurs at 36.8% (Hayek and Buzas 1997). When sites have the same number of taxa, evenness can be compared (Hayek and Buzas 1997). In the present study, LBB and DH had 20 taxa, CB and EF had 23 taxa, and L@DH and 2013 THE GREAT LAKES ENTOMOLOGIST 73

Table 6. Shannon diversity index (H´) for taxa collected from riffle habitats of 10 of 11 sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois (CI equals 95% confidence interval).

Study Siteab H´ c SE Lower CI Upper CI RB 1.07 0.04 0.99 1.15 DH 1.28 0.05 1.19 1.37 L@Rd 1.34 0.02 1.29 1.38 LBB 1.49 0.07 1.34 1.63 BB 1.46 0.03 1.40 1.53 L@DH 1.64 0.02 1.60 1.67 CB 1.79 0.03 1.73 1.86 LL@L 1.88 0.03 1.83 1.93 L@LL 1.93 0.02 1.88 1.97 EF 2.01 0.04 1.94 2.09 aSee Appendix 1 for complete names and stream order. bLL@EF was not included (see text). c {-∑pi ln pi}, p i= proportion of the ith taxon.

Table 7. Dominant taxa showing differences in evenness from riffle habitats of 10 of 11 sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Taxona

Study Sitebc Orthocladiinae Simulium spp. Chironomini Tanytarsini

RB 72.4 2.6 2.1 16.0 DH 69.6 0.8 9.6 3.1 L@Rd 21.4 53.6 5.5 6.6 LBB 62.2 2.6 3.9 3.3 BB 58.5 20.0 3.9 3.4 L@DH 39.8 13.6 30.1 5.8 CB 41.9 17.9 12.1 14.4 LL@L 31.3 28.4 16.8 6.6 L@LL 36.2 21.3 11.2 12.3 EF 22.3 30.6 19.8 6.7 aTotal number of individuals of selected taxon divided by total number of individuals for the site (e.g., Orthocladiinae, 891/1,231 = 72.4%; see Table 1). These percentages indicate amount of evenness and influence Shannon diversity index values. bSee Appendix 1 for complete names and stream order. cLL@EF was not included (see text). 74 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

LL@L had 30 taxa (Tables 1 and 2). All paired sites had different H´ values (Table 6) indicating differences in evenness (Table 7). Lusk Creek results were similar to other investigations of relatively undisturbed streams. Allan (1975) examined a Colorado stream from June to August and calculated H´ values from Surber sampling results for the EPT and Coleoptera (he excluded the Diptera) and reported values of 0.962 to 1.983. He did not note the log base used for his calculations. Initially, log base 2 was used and now the trend is to use the natural log (Magurran 2004). Wu and Legg (2007) investigated two Wyoming stream systems in summer and fall and reported H´ values of 1.33 to 2.10 using the natural log for their calculations.

The three richness estimators, first-order jackknife, Chao1, and Chao2, were compared. The first-order jackknife estimate (Manly 1997) was 74.40 (Tables 8 and 9) and predicted approximately 14 unobserved taxa (in addition to the 60 observed taxa [Tables 1 and 2]). The Chao1 estimate (Colwell and Coddington 1994) was 97.50 (Table 9) and predicted approximately 37 unobserved taxa.

The Chao2 estimate (Colwell and Coddington 1994) was 70.67 (Table 9) and predicted approximately 10 unobserved taxa. A taxonomic survey usually does not account for all taxa that are present (Chao 2005). The observed number of taxa is the lowest possible estimate of the true richness (Smith and Pontius 2006), and, therefore, the observed richness often is an underestimate (Longino et al. 2002). Methods, including first-order jackknife, Chao1, and Chao2, have been used to estimate the true richness from random samples (Colwell and Coddington 1994). These nonparametric richness estimators do not assume the frequency at which new species will be encountered (Chao 2005). However, the community distribution does influence the estimated richness (Colwell and Coddington 1994). The first-order jackknife estimate is driven by unique taxa (i.e., those found at one site only) (Heltshe and Forrester 1983). Therefore, for a given site, as the number of unique taxa increases, the pseudo-value increases. The jackknife estimate is the mean of the pseudo-values (Manly 1997). Reporting the results of a simulation experiment, Manly (1997) noted that the jackknife reduced the bias of the estimate but did not perform well in the calculation of the standard error.

Chao1 utilizes abundance information (Magurran 2004). The Chao1 estimator is calculated using taxa represented by one (singletons) or two (doubletons) specimens (Colwell and Coddington 1994). As the number of singletons increases relative to doubletons, the estimate increases in value (Chazdon et al. 1998).

Chao1 should not be used when there are major ecological differences among sites (Magurran 2004). Because Chao1 utilizes abundance information, it is sensitive to non-random distributions (Chazdon et al. 1998).

Chao2 utilizes occurrence information (Colwell and Coddington 1994, Magurran 2004) and is calculated using taxa that are found at one (unique) or two (duplicate) sites (Chazdon et al. 1998). Using seed bank data, Colwell and Coddington (1994) compared six nonparametric estimators and found that for small samples (minimum of twelve), Chao2 had a high degree of accuracy. Chazdon et al. (1998) found that Chao2 was tolerant of small sample size and moderate non-randomness. When comparing estimators, Silva and Codding- ton (1996) noted that Chao2 may be “the most practical.” As with Chao1, Chao2 should not be used when there are major ecological differences among sites (Magurran 2004).

First-order jackknife, Chao1, and Chao2 are limited because they have maximum values (Silva and Coddington 1996). For the first-order jackknife, the maximum value is approximately two times the observed number (Colwell and

Coddington 1994). For Chao1 and Chao2, the maximum value is approximately the observed number squared divided by two (Colwell and Coddington 1994). 2013 THE GREAT LAKES ENTOMOLOGIST 75 n 59 69 L@Rd 56 96 L@LL 81.31 74.99 115.90 Upper CI 58 78 LL@L LL@L 58 78 L@DH L@DH 67.49 79.10 63.35 Lower CI 59 69 EF

ab SE 3.06 9.39 2.20 60 60 CB Study Sites 58 78 BB SD 9.67 6.97 29.69 29.69 58 78 DH

a 74.40 74.40 97.50 70.67 = partial estimate total. estimators used to estimate richness from the taxa collected riffle habitats of 10 11 sites i = 3, standard deviation (SD), standard error (SE), and confidence intervals (CI). error (SE), and confidence intervals = 3, standard deviation (SD), 2 59 69 2 = 12, standard deviation (SD), error (SE), and confidence intervals (CI). RB Estimate S-j F , M , = 60, = 15 1 S = 16 and Chao F 1 L 59 69 LBB 60, 60, = = n = 10, , obs obs S )} S d )}, S-j )}, 2

F b M c /2 /2 2 2 1

c d L F 1 2 -((n-1)* +( +( S obs obs Partial Estimate Pseudo-value Jackknife Chao Chao S S {n* { See Appendix 1 for complete names and stream order. LL@EF was not included (see text). LL@EF was not included (see text). Mean of the pseudo-values (Table 8), standard deviation (SD), error (SE), and confidence intervals (CI). Total taxa (60 [see Table 2]) minus total number of unique (i.e., times partial estimate equaled zero for site). { a b c d Table 8. Partial estimate totals and pseudo-values used in jackknife calculations for taxa collected from riffle habitats of 10 11 sites in 2003 insect survey of Lusk Creek Pope County, Illinois. Table 9. Jackknife, Chao 2003 insect survey of Lusk Creek in Pope County, Illinois (CI equals 95% confidence interval). a b c d 76 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Therefore, the first-order jackknife is more affected by undersampling (Silva and Coddington 1996). The Chao and jackknife estimators have been compared to other methods and found to perform well. Using a well-known bird community, Walther and Martin (2001) compared 19 estimators and found the Chao methods were the most precise and least bias followed by the jackknife methods. In their review of numerous studies and several different estimators, Walther and Moore (2005) found the jackknife and Chao estimators generally were the most accurate. Determining richness estimates with different methods is recommended (Walther and Moore 2005). The first-order jackknife yields a conservative estimate. Therefore, if other estimators yield similar results, that would suggest “a robust estimate” (Silva and Coddington 1996). The performance of the first-order jackknife and the Chao methods have been evaluated by other investigators and their assessments reported here. In the present study, three richness estimators were calculated using a real data set. The actual performance of each richness estimate was not tested. Although confidence intervals were determined, those for the jackknife estimate are known not to perform well, and others have improved performance as sample size increases. Because the first-order jackknife and Chao estimators reflect the data, they may not always produce good estimates (Walther and Moore 2005). Therefore, the actual performance of the estimators in the present study is unknown. The difference in the number of individuals between sites is a potential source of sampling error. For the present study, the samples were standard- ized by area, which resulted in a different number of individuals per site. Most sites had between 1,000 and 3,500 individuals (Table 1). The exceptions were LBB with almost 400 individuals and L@DH with more than 4,000 individuals (Table 1). Observed richness often is correlated with sample size (Lande et al. 2000). If a correlation had been present, LBB and L@DH jackknife results would have been affected by this sampling error because the jackknife estimate is an average of pseudo-values (Table 8). However, LBB and L@DH had one and two unique taxa, respectively, which were typical values (Table 8). Chazdon et al. (1998) found that a non-random distribution or “patchiness” can be detected by comparing the number of singletons to uniques and the number of doubletons to duplicates. When they are similar, the distribution is random. For the present study, a comparison of the 15 singletons and 16 uniques (Table 1) suggested a random distribution. However, a similar comparison of the 3 doubletons and 12 duplicates (Table 1) suggested a nonrandom or patchy distribution. The edge explanation of rareness (i.e., meeting of adjacent habitats), which results in a high number of singletons, may have countered the non-randomness of the singletons compared to the uniques. Richness estimates were calculated using a small sample (10 sites), and the distribution exhibited some degree of patchiness. The Chao2 has been shown to be accurate with small samples and tolerant of moderate patchiness. Of the three estimators, it produced the most conservative estimate of unobserved taxa. In addition, it was similar to the jackknife estimate, and agreement among estimators suggests a robust result. However, given the small number of samples, these results most likely represent an underestimate.

All sites were ecologically similar, which is a requirement of Chao1. How- ever, the Chao1 estimate was larger than the other two estimators. This result may be due to the patchy distribution, as the Chao1 is known to be sensitive to patchiness. As mentioned above, the number of doubletons was small relative to the number of duplicates. This would seem to be indicative of clumping. When many individuals of all taxa are collected after extensive sampling, it would seem that the survey is complete (Coddington et al. 1996). However, 2013 THE GREAT LAKES ENTOMOLOGIST 77 rare taxa will persist even after extensive collecting (Mao and Colwell 2005). For the present study, the results represent a small number of samples using one collection method from one habitat in one season. Given the limitations of this study and the high number of rare taxa, the results indicate an incomplete survey. To date, this study is the first comprehensive investigation of the Lusk Creek spring/summer aquatic insect community and supports its recognition as a high quality and biologically signficant area. However, much work remains to be done. Similar studies should be conducted during the summer and fall months and for more than 1 year to provide a more complete picture of the richness of the Lusk Creek community. This proposed study should be conducted soon because the high quality of the area, undoubtedly, will be subjected to further anthropogenic influences.

Acknowledgments We thank the following current and former faculty, staff, and students of Southern Illinois University at Carbondale: Brooks M. Burr (Department of Zoology) for his critical review of an earlier version of the manuscript; John D. Reeve (Department of Zoology) for his advice on the statistical aspects of this project; Kevin Davie and Nathan Rahe (Morris Library Geospatial Resources) who provided assistance with mapping and figures, respectively; and William L. Muhlach (Department of Zoology) who helped the second author obtain Federal Work Study matching funds for two undergraduate student workers, Angela Defore and Kristi Magraw, who provided invaluable assistance with sample analysis. Frank M. Wilhelm and Mike Venarsky in the Limnology Laboratory; and Tom Heatherly, Scott Peterson, and Mandy Stone in the Stream Ecology Laboratory; provided technical assistance and support. We also thank R. Edward DeWalt (Illinois Natural History Survey) for identifying several of the taxa and verifying others. We are grateful to Jody Shimp (Illinois Department of Natural Resoures) for providing help in obtaining an IDNR grant (Wildlife Preservation Fund Grant #04-30W) for this project, John Taylor (United States Forest Service) for permission to sample in the Shawnee National Forest, Kris Twardowski (United States Forest Service) for mapping assistance, and Tara Kieninger (Illinois Department of Natural Resources) for Illinois Natural Area Inventory Information. We also are grateful to C. S. Bundy (Department of Entomology, Plant Pathology, and Weed Science, New Mexico State University, Las Cruces) for final changes and preparation of the figures for publication. This manuscript was part of a thesis submitted to Southern Illinois University by J. M.T. in partial fulfillment of the requirements for the M.S. degree in Zoology.

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e ) 2 10.70 11.91 18.87 21.13 41.22 44.80 2.07 4.15 4.98 8.18 110.91 (km Watershed Area = Little Bear Branch, LBB d Status Perennial Perennial Perennial Perennial Perennial Perennial Perennial Hydrologic Intermittent Intermittent Intermittent Intermittent = Ramsey Branch (See Fig. 3).

c = Little Lusk Creek at (downstream of RB = Lusk Creek at Little (downstream of /sec) 3 30.1 29.3 58.7 63.2 91.0 131.6 105.6 LL@L no flow no flow no flow no flow (m Base flow L@LL = East Fork (Little Lusk Creek tributary), b EF 0.09 0.09 0.16 0.09 0.12 0.11 0.12 0.12 0.19 0.20 0.21 (m) Channel Depth = Dog Hollow, b DH = Lusk Creek (at Eddyville Blacktop), and 4.22 5.42 7.76 5.11 5.15 7.34 3.85 6.65 10.71 10.38 13.55 (m) L@Rd Channel Width 1 1 2 2 2 2 2 3 3 4 1 = Lusk Creek at Dog Hollow (upstream of Hollow), = Copperous Branch, Order Stream CB L@DH a = Little Lusk Creek at East Fork (upstream of confluence with Fork), = Bear Branch, Study Site LBB RB DH BB CB EF L@DH LL@EF LL@L L@LL L@Rd Intermittent = no flow on day of base measurement. BB Measured during April 2005. Determined with a digital map and GIS. Measured during September 2003. Appendix 1. Selected physical properties and watershed area of sites used in 2003 insect survey Lusk Creek Pope County. a LL@EF Martha’s Woods), Lusk Creek Canyon Nature Preserve), b c d e 2013 THE GREAT LAKES ENTOMOLOGIST 85 g 0.00 0.00 1.15 0.00 0.10 0.00 0.01 0.09 0.00 0.00 0.05 0.07 Other

0.22 0.20 0.02 0.21 0.10 0.20 0.10 0.01 0.09 0.12 0.11 Water a

e 2.58 2.58 6.62 6.62 2.92 3.22 5.92 5.93 22.65 22.65 13.65 11.07 5.19 4.26 Row Crop Landcover (%) d 5.46 16.58 16.58 29.56 19.01 19.44 12.10 17.64 14.44 7.76 9.07 29.66 29.66 Grass c 78.01 46.44 91.95 67.04 69.39 83.43 63.53 89.32 87.62 76.26 79.45 Forest

Study Site LBB RB DH BB CB EF L@DH LL@EF LL@L L@LL L@Rd Includes grassland, pasture/hay, urban parklike, and herbaceous wetland. Determined with a digital map and GIS. See Appendix 1 for complete names and stream order. Includes low residential and commercial/industrial/transportation areas. Includes row crop and small grain. Includes deciduous, evergreen, mixed, and woody wetland. Includes all open water. Appendix 2. Land cover percentages of sites used in 2003 insect survey Lusk Creek Pope County, Illinois. a b c d e f g 86 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Appendix 3. Watershed drainage density, hillside slope, and main channel sinuosity values of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

DrainageDensity Main Channel Study Sitea (km/km2)b HillsideSlope%c Sinuosityd

LBB 1.47 15 1.17 RB 1.23 8 1.36 DH 1.33 6 1.22 BB 1.20 10 1.64 CB 1.87 2 1.68 EF 2.04 3 1.31 L@DH 2.01 1 1.30 LL@EF 1.46 1 1.44 LL@L 1.71 1 2.18 L@LL 1.65 3 3.36 L@Rd 1.71 1 4.96 aSee Appendix 1 for complete names and stream order. bCalculated with digital map and GIS and derived by dividing total stream system length by area. cCalculated with digital map, topographic maps, and GIS and derived by dividing elevation (rise) by run and multiplying by 100. dCalculated with digital map and GIS and derived by dividing main channel length by main channel distance. 2013 THE GREAT LAKES ENTOMOLOGIST 87

Appendix 4. Percent canopy cover and stream bed substratum composition of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Substratum (%)a Bedrock/ Canopy Study Siteb Cobble Gravel Boulder Sand Otherc (%)a

LBB 31 48 17 1 3 72 RB 53 36 9 2 0 41 DH 35 46 13 3 3 40 BB 42 36 22 0 0 40 CB 35 56 9 0 0 14 EF 43 47 8 0 2 45 L@DH 58 25 15 0 2 28 LL@EF 38 43 10 9 0 28 LL@L 54 23 16 6 1 37 L@LL 41 26 29 3 1 44 L@Rd 32 46 2 17 3 9 aMeasured during April 2005. bSee Appendix 1 for complete names and stream order. cIncludes clay, clay/sand, wood, and plant.

Appendix 5. Percent riffle/run and pool estimated visually for sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Study Sitea Riffle/run (%)b Pool (%)b LBB 58 43 RBc 28 73 DH 69 31 BB 59 41 CB 64 36 EF 85 15 L@DH 89 11 LL@EFc 46 54 LL@L 79 21 L@LL 88 13 L@Rd 90 10 aSee Appendix 1 for complete names and stream order. bMeasured during April 2005. cRiffles separated by elongated pools. 88 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Appendix 6. Water temperature during June 2003–April 2004 of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Month Jun Aug Sept Nov Jan/Feb Apr

Study Sitea Water Temperature (oC)

LBB 18.8 21.5 15.4 9.3 2.3 18.9 RB 20.3 Dry Dry Dry 1.6 14.5 DH 20.5 20.6 17.0 11.7 4.1 13.2 BB 21.6 22.7 16.7 9.3 3.4 20.9 CB 23.8 25.2 20.7 11.9 3.0 12.3 EF 20.1 20.4 16.6 7.5 2.8 14.1 L@DH 23.6 23.0 17.5 7.3 4.2 13.8 LL@EF 21.1 21.4 17.5 7.4 3.1 13.3 LL@L 23.1 23.6 17.8 10.0 1.9 12.2 L@LL 22.6 23.5 18.6 10.9 1.1 16.2 L@Rd 24.9 26.5 19.1 10.9 1.2 12.1

aSee Appendix 1 for complete names and stream order.

Appendix 7. pH during August 2003–April 2004 of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Month Study Sitea Aug Nov Feb Apr LBB 5.99 6.22 6.67 6.46 RB Dry Dry 7.82 7.30 DH 5.78 6.22 6.49 6.39 BB 6.69 7.00 7.46 6.86 CB 7.36 7.36 8.12 7.88 EF 7.06 6.97 6.69 6.63 L@DH 7.18 6.99 6.69 6.52 LL@EF 7.15 6.81 6.50 6.57 LL@L 6.73 6.85 6.98 6.46 L@LL 6.61 7.06 7.08 6.66 L@Rd 6.96 6.93 7.07 6.67

aSee Appendix 1 for complete names and stream order. 2013 THE GREAT LAKES ENTOMOLOGIST 89

Appendix 8. Conductivity during June 2003–April 2004 of sites used in 2003 insect survey of Lusk Creek in Pope County, Illinois.

Month

Jun Aug Sept Nov Jan/Feb Apr

Study Sitea Conductivity (µS/cm)

LBB 65.9 73.9 105.5 78.9 51.9 65.1 RB 472.5 Dry Dry Dry 181.7 340.0 DH 58.5 65.0 72.0 77.5 62.1 65.7 BB 109.8 148.9 123.8 159.5 90.9 136.0 CB 338.8 440.6 417.5 487.2 189.2 301.0 EF 124.3 121.5 93.4 221.2 ----b 93.0 L@DH 95.6 97.0 98.3 110.0 108.8 115.6 LL@EF 81.3 82.5 85.3 104.0 81.9 79.5 LL@L 97.7 85.6 97.6 123.2 60.8 90.1 L@LL 98.9 89.3 108.8 168.9 71.1 104.9 L@Rd 121.4 86.5 91.0 116.8 87.8 112.6 aSee Appendix 1 for complete names and stream order. bUnable to obtain reading from meter. 90 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Impact of Mecinus janthinus (Coleoptera: Curculionidae) on the growth and reproduction of Linaria dalmatica (Scrophulariaceae) Elizabeth J. Goulet1, Jennifer Thaler2, Antonio Ditommaso3, Mark Schwarzländer4 and Elson J. Shields2*

Abstract Dalmatian toadflax, Linaria dalmatica (L.) Mill. (Scrophulariaceae), a native to the eastern Mediterranean and Black Sea regions of Europe and Asia, has invaded over one million hectares in the western United States and Canada, in habitats similar to its native range. Once established, the aggressive vegetative growth of the plant allows it to invade undisturbed habitats where it can out-compete most other vegetation, placing native plant communities at risk. Biological control of L. dalmatica with Mecinus janthinus Thomson (Cole- optera: Curculionidae) has shown promise in the field. In both studies reported in this paper, the presence of insect attack reduced L. dalmatica plant growth and reduced plant reproductive potential. In a field sleeve cage study, insect- attacked stems were significantly shorter (18 cm) and had 50-70% fewer fruits and flowers than the control stems at the end of the study period. M. janthinus attacked stems showed little apical growth, fewer fruits and flowers, and lower stem biomass relative to control stems. Similar results were observed in the potted plant study where the influence of the extensive root system of the plant was eliminated. This negative impact by the insect is caused both by adult feeding in the apical portion of the plant and the physical destruction of the plant stem from larvae feeding. The decrease in the insect-attacked stem heights may also have an impact on seed dispersal from the mature reproductive structures. A combination of decreased seed production through M. janthinus biological control and poor seedling competition in the moisture limited sites common to north-central Washington State and other similarly dry habitats may negatively influenceL. dalmatica populations more than general models predict.

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Dalmatian toadflax,Linaria dalmatica (L.) Mill. (Scrophulariaceae) was introduced in the United States as an ornamental by 1894 and first planted in Canada in 1901 (Vujnovic and Wein 1997). L. dalmatica, a native to the eastern Mediterranean and Black Sea regions of Europe and Asia, has invaded over one million hectares in the western United States and Canada. The invaded habitats are similar to its native range. Its closest relatives include yellow toadflax (Li- naria vulgaris Mill.) and narrow leaved toadflax Linaria( genistifolia (L.) Mill.) also considered invasive species in North America (Sheley and Petroff 1999). Individual L. dalmatica plants are short-lived perennials and can survive up to four years (Robocker 1974), but stands can persist for long periods of time from

1Department of Biology, North Seattle Comm. College, 9600 College Way N, Seattle, WA 98103. 2Department of Entomology, Cornell University, Ithaca, NY 14853. 3Department of Crop and Soil Sciences, Cornell University, Ithaca, NY 14853. 4Department of Plant, Soil and Entomological Sciences, University of Idaho, Moscow, ID 83844. *Corresponding author: (e-mail: [email protected]) 2013 THE GREAT LAKES ENTOMOLOGIST 91 vegetative propagation and prolific seed production (Robocker 1970). Seeds can remain dormant in soil for up to 10 years under field conditions. Linaria dalmatica often invades well drained disturbed sites such as road cuts and overgrazed rangeland. Once established, the aggressive vegetative growth of the plant allows it to invade undisturbed habitats where it can out- compete most other vegetation, placing native plant communities at risk (Sheley and Petroff 1999). The plant is toxic to cattle (Vujnovic and Wein 1997) though sheep are reported to graze it without adverse effects (Sheley and Petroff 1999). Biological control of weeds is often unsuccessful because the control organism does not become established or fails to reduce the fitness of the target plant (McEvoy and Coombs 1999, Davis et al 2006). Biological control of L. dalmatica with Mecinus janthinus Thomson (Coleoptera: Curculionidae) has shown promise in the field. M. janthinus is a univoltine stem boring weevil specific to a small number of perennial Linaria species with stem diameters greater than 0.9 cm. Like its Linaria spp. host plants, M. janthinus is native to southern and central Europe and southern Russia and was first released in British Columbia, Canada in 1991 (McClay and De Clerck-Floate 2002). Monitoring of 22 release sites in British Columbia, Canada (1991-94) showed 100% establishment for the beetle and some of the sites achieved attack on 100 percent of stems after only three years (De Clerck-Floate and Harris 2002, Van Hezewijk et al. 2010). The damage to L. dalmatica stems by both adult and larval feeding has been described at both individual stem and stand levels in the field (De Clerck-Floate and Harris 2002). While M. janthinus has successfully established, its role in controlling L. dalmatica has not been documented in the literature for the northwestern United States. In many field locations, attack may only occur on a small number of stems and the large root system of L. dalmatica may play a substantial role in compensatory growth in the plant after attack. Questions arise about root reserves and their influence on insect damagedL. dalmatica stems. The purpose of the studies reported herein was to assess the impact of M. janthinus feeding on L. dalmatica growth and reproduction. In the field, single stems from a ramet were selected as the experimental unit to reduce plant-to- plant variation. A related study, utilizing a single stem from a potted rhizome, was initiated to suggest if resource sharing was evident between stems when only a single stem was attacked by the insect.

Materials and Methods Sleeve cage study. The sleeve cage study was conducted at a municipal water supply well site, 341 m in elevation near Leavenworth, Washington, USA (47º 34.33’N, 120º 40.11’W) in an area approximately one hectare in size. The study location was composed of sand, gravel and cobbles and was a flat, highly mechanically disturbed area. Native plants were sparse and limited to some hardy shrubs. Dalmatian toadflax (L. dalmatica) was present throughout the site and M. janthinus was absent from the site. Thirty-three pairs of stems were selected for the cage study on 1 and 3 June 2005. Each pair of stems was selected from the same ramet of stems and was similar in height (± 3 cm), composed of approximately equal stem diameters and numbers of flowers. Stem breakage resulted in eight of the replicates being removed from the study, leaving 25 remaining pairs. The likelihood that the paired stems were from the same genotype was very high since the stems were in close proximity to each other and L. dalmatica is a rhizomatous species. Before the stems were caged, stem height, number of flowers per stem, and number of fruits per stem were recorded. Sleeve cages, approximately 90 cm long and 30 cm wide, were placed over each stem and held around the stem with paper-covered wire twist ties. 92 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

All of the insects used in this study were collected at the USDA APHIS PPQ insectary near Danville, Washington (48º 59.31’N, 118º 30.29’W), two days before being placed in the cages in Leavenworth, a distance of 140 miles by direct route. Insects were stored in cardboard containers with L. dalmatica stems and moist sponge cubes in a cooler with ice until they were placed on the study plants. For each pair of stems, one stem was randomly selected to receive 12 adult M. janthinus in the sleeve cage and the second stem served as the control. All stems were caged for nine days. Cages and insects were then removed from the stems, with both stems in each set receiving the same amount of handling. An additional 14 stems of similar size and height, selected from existing study ramets, served as uncaged controls to test for cage effects. These control stems were handled similarly to their caged counterparts. When the study was initiated, stem height, number of flowers per stem, and number of fruits per stem were recorded for the control stems. Once cages and insects were removed, stem height, number of flowers per stem, and number of fruits per stem were recorded for all of the stems in the study, including the uncaged controls. All stems were allowed to grow without cages for 46-48 d. On July 28-29, 2005, all stems were cut, measured (stem height, number of flowers, number of fruits, number of branches, and the stem diameter at the base), and dissected to determine the number of M. janthinus larvae. Once M. janthinus larvae were counted and removed, stems were oven- dried for 48 h and the weight of each stem including fruits was measured to the closest 0.01 g. Potted plant study. Linaria dalmatica rhizomes were selected from four locations in Chelan and Douglas County, Washington, USA in mid-March 2006. At each location, eight to 10 overwintering rosettes were collected leaving 30-60 cm lateral roots. Rosettes were packed in soil, wrapped in plastic, and kept in a cold dry storage area until the initiation of the study. Sections of rhizomes with roots attached from each location were cut to 0.5 cm lengths with one visible stem bud. The rhizome sections were placed in large aluminum roasting pans, covered with sterilized potting soil and watered daily until emerging shoots had grown to at least 5 cm height. Sprouted rhizomes were individually transferred to 20 cm by 14 cm plastic pots containing local soil from Chelan County. Pots were watered daily, placed in two rows outside the west facing side of a building where plants received midday to mid-afternoon sun, and any new shoots were clipped, restricting each pot to only one stem. The locations of the pots were rotated every 2 days to minimize the effect of pot location on results. Plants were grown in these pots for two years before the initiation of the study. In late June 2008, 18 stems of similar vigor with heights ranging from 27 - 142 cm were selected for the study. Pots were divided into three groups of six plants each with similar stem heights. An additional 10 stems of similar height and vigor were selected as untreated controls. Data collected from each stem included height, number of branches and number of leaves in the top 10 cm of the stem. All stems except controls were covered with net sleeve cages, held away from the plant with wire. M. janthinus adults used for the study were collected in late June 2008 at the USDA APHIS PPQ insectary near Danville, Washington. Stems in each of the three groups randomly received between two to eight pairs of insects with each stem within a group receiving a different number of insect pairs. Each stem was labeled and the number of insect pairs introduced into the cage was recorded. The study design provided six different numbers of insect pairs on stems, replicated three times. After six days, cages and insects were removed from the stems. Stems were then allowed to grow cage free for the rest of the growing season and watered daily. Although no cages were placed over the control stems, the stems were handled in a similar manner as the caged plants to equalize the negative impact of handling across all treatments (Niesenbaum et al. 2006). 2013 THE GREAT LAKES ENTOMOLOGIST 93 In mid-November 2008 all potted plants were clipped at the soil surface level and stems dissected to determine the number of live and dead M. janthinus adults found in each stem. The remainder of the underground stem was clipped from the root and added to the aboveground stem for biomass measurements. Data recorded included stem height, number of branches, number of fruits and flowers, and the number of leaves in the top 10 cm of the main stem. Stems and roots were then rinsed with water, bagged individually and oven-dried to determine biomass.

Data analysis Sleeve cage study. Comparisons among L. dalmatica stem heights and the numbers of fruits and flowers were calculated across treatments using pre- treatment and post-treatment data. The general model, Dependent variable = treatment type + block size + significant interactions,was used for the analysis. Stems from each ramet served as a block with either two stems (insect attacked and cage only) or three stems (an additional uncaged control). The analysis of numbers of fruits and flowers included blocks in which stems from all three treatments had at least one flower or fruit post treatment. Mixed model analyses were used on both a comparison of dry stem biomass (g) and mean dry fruit weight. The relationship between number of larvae in stems and the growth of M. janthinus-attacked stems was evaluated using regression analysis. The relationship between number of larvae in stems and the change in number of fruits and flowers in insect-attacked stems was evaluated using the same regression model as for stem height. Natural log (ln) transformation was used for stem height, number of fruits and flowers, and biomass values to normalize data. Fruit weight and numbers of insects were normally distributed and were not transformed for analyses. A Bonferroni adjustment was made to control for error rate in the linear mixed model analysis. All statistical analyses were conducted using SAS version 9.1 (SAS Institute 2006) and P values greater than 0.05 were considered not significant. Potted plant study. A comparison between attacked and control L. dalmatica stem heights was calculated across treatments and across time with a linear mixed model with treatment and time as fixed effects in the model. A Bonferroni adjustment was used to control for error rate. T-tests were used to examine differences in the number of fruits and flowers, root biomass, and stem biomass data. The relationship between number of larvae and root biomass as well as stem height was assessed with regression analyses. Stem heights and above ground biomass were converted to natural log values to normalize data for analysis and square root transformation was used to normalize values in the fruits and flowers analysis. All statistical analyses were conducted using SAS version 9.1 (SAS Institute 2006) and P values greater than 0.05 were considered not significant.

Results Sleeve cage study. Comparison between the caged controls and uncaged controls showed no cage effect on stem height in the experiment (n=13). Unlike control stems, the M. janthinus attacked stems did not grow over the duration of the study and were 11% shorter than both the caged control (t = -6.37, df = 24, P = <0.001) and uncaged control stems (t = -4.45, df = 24, P = <0.001) after treatment (Table 1). There was no height difference in the insect-attacked stems before and after treatment (t = 0.30, df = 24, P = 1.00). In addition, insect-attacked stems weighed 17% less than control stems (t = 5.71, df = 21, P = 0.0079) (Table 2). While initially there were no differences in flower and 94 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 fruit numbers across the treatments, significant differences were recorded at the end of the study. Post treatment comparisons indicated that insect-attacked stems had fewer flowers and fruits than both the caged stems (t = -3.35, df = 14, P = 0.019) and control stems (t = -3.31, df = 14, P = 0.021) (Fig 1, Table 3). However, there were no differences in individual fruit weight between any of the treatments. Comparison between the caged controls and uncaged controls showed no cage effect on flower and fruit numbers in the experiment. The number of larvae ranged from four to 37 per stem. No relationship was found between change in stem height and the number of larvae in a stem (F = 1.01, r 2 = 0.096, P = 0.382,). In addition, the number of larvae per stem did not predict the change in numbers of fruits and flowers per stem (F = 1.61, r2 = 0.244, P = 0.247). Potted plant study. The untreated control stems grew on average an additional 18.7 cm through the season; a significant height increase compared with the pre-treatment measurements (t = 4.78, df = 5, P = 0.0008). In contrast, the height of the M. janthinus attacked stems remained unchanged throughout the growing season (t = 0.74, df = 15, P = 1.00) (Table 4). When the study was initiated, no stems had flowers. At the end of the study, most stems (treated or untreated) did not produce flowers or fruits. They were, however, produced on fiveM. janthinus-attacked stems (9.8 ± 2.3 per stem) and seven untreated control stems (20.4 ± 4.2 per stem) and the two groups were significantly different from each other t( = -2.17, df = 4, P = 0.0551). At the end of the study, the biomass of the untreated control stems (2.03 ± 0.293 g) was significantly higher than the insect attacked stems (1.39 ± 0.210 g) (t = 2.08, df = 9, P = 0.0571). There was no difference in root biomass between the two groups. A correlation between the number of insect pairs placed on each treated plant and the change in stem height through time was noted. The model, using the difference in stem height as the dependent variable and the starting height of the stem as a covariant, indicated differences were significant (F = 4.78, R2 = 0.42, P = 0.0279). Parameter estimates showed that numbers of pairs of insects was significant t( = -2.90, df = 5, P= 0.0123) and the covariate was not significant (t = 0.43, P = 0.675). There was no relationship, however, between the number of larvae in the stems and number of fruits and flowers, difference in stem height, or stem biomass.

Discussion In both studies, the presence of insect attack reduced L. dalmatica plant growth and reduced plant reproductive potential. In the field sleeve cage study, insect-attacked stems were significantly shorter (18 cm) and had 50-70 percent fewer fruits and flowers than the control stems at the end of the study period. M. janthinus attacked stems showed little apical growth, fewer fruits and flowers, and lower stem biomass relative to control stems. Similar results were observed in the pot study where the influence of the extensive root system of the plant was eliminated. This negative impact by the insect is caused both by adult feeding in the apical portion of the plant and the physical destruction of the plant stem from larvae feeding. It is interesting to note that there appears to be no relationship between the number of M. janthinus larvae in the stem and their impact on stem height, though work by Van Hezewijk et al. (2010) found a strong negative correlation between M. janthinus density and stalk length across years at study sites in British Columbia. No correlation between insect density and stalk density was determined in their work. In addition, our data suggest that the number of M. janthinus larvae in stems appears to be a poor predictor of the degree of 2013 THE GREAT LAKES ENTOMOLOGIST 95

Table 1. Mean height (± SE) of uncaged control, caged control, and M. janthinus-attacked L. dalmatica stems pre and post treatment. Different letters indicate significant difference at the α = 0.05 level.

Treatment Time Number of Stem height stems (cm) ± se

Uncaged control Pre treatment 13 72.5 ± 9.1 a

Post treatment 13 80.9 ± 11.7 b

Caged control Pre treatment 25 71.3 ± 5.5 a

Post treatment 25 79.9 ± 6.9 b

Insect-attacked Pre treatment 25 71.1 ± 5.6 a

Post treatment 25 71.0 ± 5.6 a

Table 2. Mean biomass (g) (± SE) of M. janthinus-attacked, caged control, and uncaged control L. dalmatica stems post treatment. Different letters indicate significant difference at α = 0.05 level.

Treatment Number of stems Stem biomass (g) ± se Insect-attacked 22 3.26 ± 0.52 a

Caged control 22 3.93 ± 0.63 b

Uncaged control 10 3.49 ± 0.98 ab

Figure 1. Mean number of fruits and flowers (± SE) onM. janthinus-attacked, unattacked and control (uncaged) L. dalmatica stems. N = 15 stems for insect attacked and caged control. N =6 stems for uncaged control. Different letters indicate significant difference at α = 0.05 level. 96 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 3. Mean number of fruits and flowers (± SE) onM. janthinus-attacked, caged control, and uncaged control L. dalmatica stems pre and post treatment. Different letters indicate significant differences at α = 0.05 level.

Number of Number of fruits and Treatment Time stems flowers ± se

Insect-attacked Pre treatment 15 4.1 ± 1.54 a Post treatment 15 7.7 ± 2.14 a Caged control Pre treatment 15 3.7 ± 1.49 a Post treatment 15 17.2 ± 3.15 b Uncaged Pre treatment 6 7.3 ± 2.65 control a Post treatment 6 24.2 ± 4.54 b

Table 4. Mean height (cm) (± SE) of L. dalmatica potted plant stems before and after M. janthinus attack. Different letters indicate significant difference at α = 0.05 level.

Number of Group Time Height (cm) ± SE stems Control Pre treatment 6 28.0 ± 1.53 a

Post treatment 6 46.7 ± 5.02 b

Insect-attacked Pre treatment 16 31.6 ± 1.84 a

Post treatment 16 33.3 ± 1.97 a decrease in fruit and flower number in stems that are attacked. These data indicate that the impact of the insect is more closely related to their presence/ absence than the actual number of insects present on plants. No relationship could be discerned between the numbers of larvae in the stem and changes in plant growth or reproductive capacity. However, Jeanneret and Schroeder (1991, 1992) reported that larval mining in high density or outbreak populations of M. janthinus caused premature stem wilting and suppression of flower formation, and consequently, reduction in seed production. Results from the pot study, however, suggest that the number of M. janthinus adult insects on the stem does correlate with the negative impact of the beetle on change in stem height. This relationship can be partially explained by the impact of adult feeding in the apical portion of the plant. Greater damage to the apical meristem from a larger number of adult insects feeding would be expected to have a more profound effect on the ability of the host plant to grow after damage. Beetles reduced flowering both when root reserves were available and when they were not. In both the potted plants and field stem study, attacked plants had comparable numbers of flowers (9.8 average in potted plants and 7.7 in field stems). In the pot study, no differences were noted in the root biomass by treatment comparisons. Since the pot study was a single year study, these results may not reflect the cumulative impact of the insect on root biomass in 2013 THE GREAT LAKES ENTOMOLOGIST 97 situations where the insect has attacked the plant for several seasons, resulting in reduced growth for several seasons. The lack of difference in individual reproductive flower/fruit/seed weights between the insect-attacked stems and the controls suggests that the negative effect of M. janthinus feeding on the reproductive capacity of the plant may be due to decreased flower and fruit production but not due to an actual decrease in quality of the fruit and seeds. A study comparing seed numbers from fruits collected from both attacked and unattacked stems could further clarify whether there is a decrease in quality, or only quantity, of reproductive structures as a result of M. janthinus damage. The decrease in the insect-attacked stem heights may also have an impact on seed dispersal from the mature reproductive structures. Some biological control models suggest that as much as 95% or more of seeds need to be destroyed in some invasive plants if plant density is to be reduced (Myers and Bazely 2003). In addition, L. dalmatica seedlings are known to be poor competitors for moisture (Sheley and Petroff 1999). A combination of decreased seed production through M. janthinus biological control coupled with poor competition of seedlings in the moisture limited sites common to north-central Washington State and other similarly dry habitats may negatively influenceL. dalmatica populations more than general models predict.

Acknowledgments We thank Tracy Valentine of City of Leavenworth Water Plant Division, Larry Skillestad of USDA APHIS, Robert Fischer of US Army Corps of Engineers, and Lorraine Goulet for their valuable assistance on this project.

Literature Cited Davis, A. S., D. A. Landis, B. Nuzzo, B. Blossey, E. Gerber, and H. L. Hinz. 2006. Demographic models inform selection of biocontrol agents for garlic mustard (Alliaria petiolata). Ecological Applications 16: 2399-2410. De Clerck-Floate, R. A., and P. Harris. 2002. Linaria dalmatica (L.) Miller, Dal- matian toadflax (Scrophulariaceae), pp 368-374. In P. G. Mason and J. T. Hubner (eds.) Biological control programmes in Canada, 1981-2000. CAB International, Wallingford, United Kingdom. Jeanneret, P., and D. Schroeder. 1991. Final Report: Mecinus janthinus Germar (Col.: Curculionidae): a candidate for biological control of Dalmatian and yellow toadflax in North America. International Institute of Biological Control, European Station, Delemont, Switzerland. Jeanneret, P., and D. Schroeder. 1992. Biology and host specificity of Mecinus janthinus Germar (Col.: Curculionidae), a candidate for the biological control of yellow and Dalmatian toadflax, Linaria vulgaris (L.) Mill. and Linaria dalmatica (L.) Mill. (Scrophulariaceae) in North America. Biocontrol Science and Technology. 2: 25-34. McClay, A., and R. A. De Clerck-Floate. 2002. Linaria vulgaris Miller, Yellow toadflax (Scrophulariaceae), pp. 375-382. In P. G. Mason and J. T. Hubner (eds.) Biological control programmes in Canada, 1981-2000. CAB International, Wallingford, United Kingdom. McEvoy, P. B., and E. M. Coombs. 1999. Biological control of plant invaders: Regional patterns, field experiments and structured population models. Ecological Applica- tions 9: 387-401. Myers, J., and D. Brazely. 2003. Ecology and control of introduced plants. Cambridge University Press, Cambridge, United Kingdom. 98 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Niesenbaum, R., J. Cahill, and C. Ingersoll. 2006. Light, wind and touch influence leaf chemistry and rates of herbivory of Apocynum cannabinum (Apocynaceae). International Journal of Plant Sciences. 167: 969-978. Robocker, W. 1970. Seed characteristics and seedling emergence of Dalmatian toadflax. Weed Science. 18: 720-725. Robocker, W. 1974. Life history, ecology and control of Dalmatian toadflax. Washington Agricultural Experiment Station. Technical Bulletin 79. Pullman, Washington, USA. SAS Institute. 2006. SAS/STAT user’s guide, version 9.1. SAS Institute, Cary, NC. Sheley, R. L., and J. K. Petroff, (eds.) 1999. Biology and management of noxious rangeland weeds. Oregon State University Press, Corvallis, OR. Van Hezewijk, B. H., Bourchier, R. S., and De Clerck-Floate, R. A. 2010. Regional- scale impact of the weed biocontrol agent Mecinus janthinus on Dalmatian toadflax (Linaria dalmatica). Biological Control. 55: 197-202. Vujnovic, K., and R. W. Wein. 1997. The biology of Canadian weeds. 106. Linaria dalmatica (L.) Mill. Canadian Journal of Plant Science. 77: 483-491. 2013 THE GREAT LAKES ENTOMOLOGIST 99 First Report of the Adventive Species Sitochroa palealis (Lepidoptera: Crambidae) in Pennsylvania and its Attraction to the Sex Pheromone of the European Corn Borer, Ostrinia nubilalis (Lepidoptera: Crambidae) Neelendra K. Joshi1, 2*, David J. Biddinger1,2, Shelby Fleischer 2, Steven Passoa3

Abstract Sitochroa palealis (Denis and Schiffermüller, 1775) (Lepidoptera: Crambi- dae), a newly detected crambid moth in the United States, is found for the first time in orchard and row crop agroecosystems in Adams and Centre counties, Pennsylvania, during 2011. In Adams County, S. palealis male and female adults were net collected from flowers and found in white pan traps used to sample bee populations near apple orchards, while in Centre County adults were found in wire-cone traps baited with the sex pheromone of the European corn borer (E-strain), Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae). Additional records of S. palealis from four Ohio counties (Marion, Wayne, Franklin, and Delaware) are given. A brief discussion on the current economic importance of S. palealis in the United States is provided, and its importance as a non-target in European corn borer surveys is highlighted.

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Sitochroa palealis (Denis and Schiffermüller, 1775) (Lepidoptera: Cram- bidae) is known as the carrot seed moth (Beadle and Leckie 2012). Larvae feed on various weed and crop species belonging to Apiaceae (Umbelliferae) (Gaedike 1980). In the Old World, S. palealis has been reported from several Asian (Park 1979), European (Karsholt and Razowski 1996, Asselbergs et al. 2008, Shodotova 2008,) and North African (Balachowsky 1972) countries. The United States distribution includes several midwestern states (Illinois, Indiana, Michigan and Wisconsin) based on collections taken as early as 2002 (Passoa et al. 2008). There are currently records for southeastern Canada (Jean Francois Landry, pers. comm.) and the northeastern United States (Beadle and Leckie 2012), including New York (R. Hoebeke pers. comm.). Here we report the first detection of S. palealis in orchard and row crop agroecosystems of Pennsylvania, and the first information about its capture in sex pheromone traps. Unconfirmed photographs ofS. palealis from Allegheny County, Pennsylvania can be found on-line (Bugguide 2012). We also report on some distribution records from central Ohio. Sitochroa palealis Discovery and Identification. Adults of S. palealis (Fig. 1) were found in several locations in Adams and Centre counties, Penn- sylvania during 2011. In Adams County, S. palealis adults were found in 16 oz Solo® white plastic bowls (Solo Cup Company, Lake Forest, Illinois) used in water

1Pennsylvania State University, Fruit Research & Extension Center, Entomology, 290 University Dr, Biglerville, PA 17307. 2Pennsylvania State University, Department of Entomology, 501 ASI Building, University Park, PA 16802. 3United States Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine, Forest Service Northern Research Station and The Ohio State University, 1315 Kinnear Road, Columbus, Ohio, 54321 *Corresponding author: (e-mail: [email protected]). 100 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 1. S. palealis (mounted specimen) collected from Centre County, Pennsylvania during 2011. pan trap monitoring of bee species near apple orchards; two specimens were collected in early August at the Pennsylvania State University Fruit Research and Extension Center near Biglerville, PA. We subsequently collected (by net) four additional adults feeding on the flowers of wild bergamot Monarda( fistulosa L.) and wild carrot (Daucus carota L.). Moths were identified to species by D. J. Biddinger (Pennsylvania State University) using Passoa et al. (2008) and on-line resources (North American Moth Photographers Group 2012). Specimens were forwarded to the Pennsylvania Department of Agriculture, and were confirmed as S. palealis (APEPA 112493511002, M. Alma Solis). In Centre County, S. palealis adults were found in Hartstack wire cone traps (Hartstack et al. 1979, also see UKY 2012 for trap design). Four traps were placed along the edges of fields and sweet corn plots in Rock Springs, PA and monitored weekly from 4 June to 3 September, 2011. Traps were baited with sex pheromone lures from Hercon Environmental (Emigsville, PA) designed to capture European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Cram- bidae), as part of the PestWatch regional monitoring program (Fleischer et al. 2007). In Pennsylvania, and much of the northeast, two pheromone races of European corn borer exist in sympatry (O’Rourke et al. 2010): a Z-race where the sex pheromone is comprised of a 97:3 ratio of cis- to trans-11-tetradecenyl acetate, and an E-race, where the pheromone is a 1:99 ratio of these isomers (Klun et al. 1973). Two traps were baited for the E- and Z- strain of European 2013 THE GREAT LAKES ENTOMOLOGIST 101 corn borer, respectively. We only captured S. palealis in traps baited with the E-strain lures (E isomer of the sex-pheromone component, which is 99 % E and 1% Z isomers of 11-tetradecenyl acetate). The first adults were recorded in the trap captures during the week of 5 July 2011(Fig. 2). Emergence pattern of S. palealis (in terms of weekly adult captures in wire cone traps) is illustrated in Fig. 2. All specimens were identified to species, and voucher specimens are in the laboratory of S. J. Fleischer. It is interesting to note that the adults of S. palealis were capatured in traps baited with a blend of sex pheromone that is currently used to attract adult males of a particular ecotype of European corn borer. Both species are in the tribe Pyraustini (Crambidae: Pyrautinae), so this could be due to some similarities in the constituents of sex pheromone components of S. palealis and E-strain of European corn borer. With regard to Ohio, the earliest records include larvae from the heads of Daucus carota at the Marion County campus of the Ohio State University during September 2008. A series of adults was collected by R. Downer from July to the first of August at lights in Wooster (Wayne County) during 2009. Recently, adults were collected at lights or on clover at Powell (Delaware County) during early July of 2011 and 2012. Attempts to find larvae on Daucus carota at The Ohio State University main campus in Columbus, Ohio (Franklin County) were negative from 2002-2011. In 2012, larvae were collected there in late July. All the above specimens were collected by S. Passoa, unless otherwise indicated, and are in his collection. It is unlikely that S. palealis appeared first in central Ohio. Unfortunately, there are no specimens of S. palealis from Ohio in the Charles A. Triplehorn collection at the Ohio State University before 2008; thus it is unclear if the spread was from the west (Indiana), north (Michigan) or even the east (New York or Pennsylvania). Passoa et al. (2008) speculated that S. palealis may have entered North America through ports-of-entry along the Great Lakes. This hypothesis could change if specimens collected before 2002 exist from other regions.

Figure 2. Cumulative weekly adult captures of S. palealis in wire cone traps baited with sex pheromone of the E-strain of European corn borer in Centre County, Pennsyl- vania during 2011. 102 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Economic Importance and Management Implications. Because S. palealis is a newly detected immigrant species in the United States, our knowledge about its life history in local climatic conditions is limited. Larval stages of S. palealis feed on seeds of different Apiaceae. Thus, it could be a pest if carrots are grown for seeds or breeding purposes. In Europe, it destroyed carrot seeds completely while feeding on seed-heads (Balachowsky 1972). Although established in the United States for at least 10 years, S. palealis has not yet damaged crops to that extent in any parts of the known range. However, economic damage from immigrant insects is not always immediately apparent, and can change over time. One recent moth example is Noctua pronuba (L.); damage to lawns and crops were not reported until recently (S. Passoa, unpublished data). In contrast to its potential pest status, Passoa et al. (2008) speculated that larval populations could help control introduced weeds belonging to Apiaceae. This was indeed the case at Lakeshore State Park, Wisconsin, during early August 2011 (S. Borkin, pers. comm.). Larvae ate the flowers, leaves and even the stems of Daucus carota. Observed larval damage in Ohio and Pennsylvania has not yet been as severe. A third component to the importance of S. palealis is its non-target status in European corn borer traps. Besides our records, there is a series of specimens at Cornell University also taken from European corn borer pheromone traps in various counties of New York (E. R. Hoebeke, pers. comm.). We expect continued survey of the European corn borer in the northeastern United States will help clarify the true distribution of S. palealis, and it will continue to attract attention as it spreads throughout the region. This moth seems to have a complicated impact on several ecosystems, and can be collected in a variety of ways including larval surveys, and adults at lights or in pheromone traps. Many habitats are suitable (agricultural, urban areas, state parks and preserves); the only requirement seems to be presence of a hostplant.

Acknowledgments The authors would like to thank Nick Sloff for photography, Amanda Ritz and Karim Hemady for help in summer fieldwork. Roger Downer, Jean Francois Landry and E. Richard Hoebeke provided information based on their experience. Susan Borkin called our attention to the Wisconsin outbreak. Ac- cess to the Charles A. Triplehorn insect collection at The Ohio State University was appreciated.

Literature Cited Asselbergs, J. E. F., A. Seguna, and P. Sammut. 2008. Recent records of Pyraloidea species new to Malta, including two species new to the European fauna (Lepidoptera: Pyraloidea). SHILAP Revista de Lepidopterología 36: 461-475. Beadle, D., and S. Leckie. 2012. Peterson Field Guide to Moths of Northeastern North America. Houghton Mifflin Harcourt Publishing Company, Boston. 640 pp. Balachowsky, A. S. 1972. Entomologie Appliquée a l’Agriculture. Tome II. Lépidoptères. Masson et Cie. Paris, France. 575 pp. Bugguide. 2012. Identification, images, and information for insects, spiders and their kin for the United States and Canada. Available from http://bugguide.net/node/ view/133216/bgimage (accessed 12 December 2011). Fleischer, S., G. Payne, T. Kuhar, A. Herbert, S. Malone Jr., J. Whalen, G. Dively, D. Johnson, J. A. Hebberger, J. Ingerson-Mahar, D. Miller, and S. Isard. 2007. Helicoverpa zea trends from the Northeast: Suggestions towards collaborative mapping of migration and pyrethroid susceptibility. Available from Plant Health Progress doi:10.1094/PHP-2007-0719-03-RV. 2013 THE GREAT LAKES ENTOMOLOGIST 103

Gaedike, R. 1980. Beiträge zur insektenfauna der DDR: Lepidoptera-Pyraustinae. Be- itrage zür Entomologie 30: 41–120. Hartstack, A. W., J. A. Witz, and D. R. Buck. 1979. Moth traps for the tobacco bud- worm. Journal of Economic Entomology 72: 519-522. Karsholt, O., and J. Razowski. 1996. The Lepidoptera of Europe. A Distributional Checklist. Stenstrup, Denmark Apollo Books. 380 pp. Klun, J. A., O. L. Chapman, K. C. Mattes, P. W. Wojtkowski, M. Beroza, and P. E. Sonnet. 1973. Insect sex pheromones: minor amount of opposite geometrical isomer critical to attraction. Science 181: 661-663. North America Moth Photographers Group. 2012. North Americ moth photographers group. Available from http://mothphotographersgroup.msstate.edu/pinned. php?plate=12.1&sort=h (accessed 12 December 2011). O’Rourke, M. E., T. W. Sappington, and S. J. Fleischer. 2010. Managing resistance to Bt crops in a genetically variable insect herbivore, Ostrinia nubilalis. Ecological Applications 20: 1228-1236. Park, K. T. 1979. Catalogue of the Pyralidae of Korea (Lepidoptera). I. Evergestiinae and Pyraustinae. The Korean Journal of Plant Protection 18 (2):89–100. Passoa, S., G. Balogh, and M. A. Solis. 2008. Sitochroa palealis: a Palearctic pyraustine moth (Pyraloidea: Crambidae) newly introduced to North America. Proceedings of the Entomological Society of Washington. 110: 504-515. Shodotova, A. A. 2008. Pyralid moths (Lepidoptera, Pyraloidea) of Buryatia: Family Pyraustidae. Entomological Review 88: 543-557. University of Kentucky (UKY). 2012. Plans and parts list: “Texas” style cone trap for monitoring certain insect pests. Available from www.uky.edu/Ag/Entomology/ entfacts/misc/ef010.htm (accessed 16 May 2012). 104 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Inventory of Aquatic and Semiaquatic Coleoptera from the Grand Portage Indian Reservation, Cook County, Minnesota David B. MacLean1

Abstract Collections of aquatic invertebrates from the Grand Portage Indian Res- ervation (Cook County, Minnesota) during 2001 – 2012 resulted in 9 families, 43 genera and 112 species of aquatic and semiaquatic Coleoptera. The Dytisci- dae had the most species (53), followed by Hydrophilidae (20), Gyrinidae (14), Haliplidae (8), Chrysomelidae (7), Elmidae (3) and Curculionidae (5). The families Helodidae and Heteroceridae were each represented by a single species. Seventy seven percent of species were considered rare or uncommon (1 - 10 records), twenty percent common (11 - 100 records) and only three percent abundant (more than 100 records). Sixty nine percent of species were collected at only 1 or 2 habitat types, twenty two percent at 3 - 5 and only nine percent at 6 - 10 habitats types. Haliplus leopardus Roberts (Halipidae), Hydrocolus stagnalis (G. and H.) (Dytiscidae), and Hygrotus falli (Wallis) (Dytiscidae) were recorded for the first time from Minnesota. The Dytiscids Acilius mediatus (Say) ( Dytiscus alaskanus J. Balfour-Browne, Graphoderus fasciollis (Harris), Hydaticus piceus LeConte and Neoscutopterus hornii (Crotch) were recorded for the first time from the northeastern portion of the state.

______

Each year from 1999 through the early fall of 2012, members of the Grand Portage Band Environmental Division at the Grand Portage Indian Reservation, Cook County, Minnesota have sampled invertebrates to assess the environmental quality of aquatic and semiaquatic habitats. This paper presents the results of aquatic and semiaquatic Coleoptera collected from 42 sites from 2001 to the fall of 2012. The Grand Portage Indian Reservation (GPIR) occupies the extreme northeastern tip of Cook County, Minnesota and borders Ontario, Canada. The Reservation lies entirely within the North Shore Highlands of the Northern Coniferous Forest (Albert 1994, Marschner 1974) and includes many aquatic habitats such as numerous small lakes, streams, spring-fed ponds, wetlands and the Pigeon River that borders Ontario. Aquatic Coleoptera are important members of the predator, herbivore and collector feeding groups in aquatic food webs (Merritt and Cummins 1996). Aquatic habitats often support many species of Dytiscidae, Gyrinidae, Haliplidae and Hydrophilidae. Species of Elmidae, Helodidae, Hydraenidae, Noteridae and Ptilodactylidae (Larson 1987) are lotic and not well surveyed while Chrysomelidae are mostly terrestrial. Analyses of dytiscid communities have shown that repeatable assemblages correlate reasonably well with both lotic and lentic habitat characteristics (Larson et al. 2000). Four hundred and eighty five species of aquatic and semiaquatic beetles have been recorded from Ontario (Larson 1987), 242 (and possibly 118 others) from Michigan (Bright 2006), at least 357 from Wisconsin (Hilsenhoff 1995) and at least 250 from Minnesota, including the northeastern part of the state,

1Professor emeritus, Youngstown State University. Current address: 76 Walter Rd. Grand Marais, MN 55604. 2013 THE GREAT LAKES ENTOMOLOGIST 105 (Smetana 1978, 1988; Daussin 1979; Ferkinhoff and Gundersen 1983; Gundersen and Otremba 1988; Gundersen and Christopherson 1990). The objective of this paper is to provide an inventory of aquatic and semiaquatic beetles from the extreme northeastern region of the state, an area that has been little studied.

Methods Twenty eight sites were sampled in only one year, 5 in two years, 7 in three years, 4 in four years, 1 in five years, 2 in six years, and 1 in nine years. Sampling methods included the use of dip nets each year, bottle traps in 2001 and 2002, pit fall traps in 2003 and terrestrial light traps in 2001, 2002, 2003 and 2004. Most sites were sampled in two or more years but on different dates each year. Ten sites were sampled in 2001, 9 in 2002, 4 in 2003 (8 collections on 2 different dates), 12 in 2004, 8 in 2005, 6 in 2006, 2007, 2008, 7 in 2009, 6 in 2010 and 2011 and 5 in 2012. As most sites were not sampled each year and different sampling methods were employed, an extensive statistical analysis of the data was not appropriate. All aquatic habitat types were sampled by dip nets at all sites and combined for a total of at least 300 invertebrates whenever possible. Beetles and other invertebrates from dip net samples and bottle traps were placed in 70% ethanol, sorted in the laboratory and either pinned, placed on points or preserved in 70% ethanol. After identification to species, voucher specimens were deposited in the invertebrate collection of the Grand Portage Environmental Division, Grand Portage Band and the author’s collection. Some species identifications were checked by Dr. Ralph Gundersen, Professor Emeri- tus, Saint Cloud State University, Ethan Bright, the University of Michigan and Paul Tinerella of the University of Minnesota. A number of sites had been impacted by hydrological alterations such as culverts, proximity to roads and land use history (logging and homesteading) and invasive plants (Andrew Schmidt, Wetland Specialist, Grand Portage Envi- ronmental Division, pers. comm.). Each site was classified by the Band as one or more of the following habitat types: 1) coniferous bog, 2) fen, 3) poor fen, 4) marsh, 5) sedge meadow, 6) clear cold - moderately stained lake, 7) moderately stained to dystrophic lake, 8) shallow lake, 9) spring-fed lake, 10) beaver pond, 11) vernal pool/pond, 12) impoundment some with a beaver dam, 13) black spruce/balsam fir swamp, 14) shaded, clear flowing stream, 15) slowly moving, stained creek with beaver dams, 16) slowly flowing river with dense vegetation along margin, 17) small creek with rocky bottom, 18) medium-large river, or 19) shallow Lake Superior bay. Sites often had more than one habitat type. Sites sampled each year and the method(s) were determined by Margaret Watkins, Water Quality Specialist.

Results One hundred twelve species of aquatic and semiaquatic Coleoptera identified from the GPIR during the period 2001-2012, along with the collection method, months collected, numbers of records, habitat type and regional abundance, are listed in Table 1. Habitat type refers to those listed in the Methods section and regional abundance is based on published records from Minnesota and neighboring states and provinces. Haliplus leopardus Roberts (Halipidae), Hydrocolus stagnalis (G. and H.) (Dytiscidae) and Hygrotus falli (Dytiscidae) were recorded for the first time from Minnesota. Drought conditions that resulted in low water levels reduced the numbers of specimens collected in some years, e.g., 2007. The use of bottle traps only in 2001 and 2002 likely underestimated the number of species and individuals of Dytiscus. The Dytiscidae were represented by the most species (53) followed by Hydrophilidae (20), Gyrinidae (14), Haliplidae (8), Chrysomelidae (7), Elmidae (3) and Curculionidae (5). The families Helodidae and Heteroceridae were represented by only a single species 106 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Families, genera and species of aquatic and semiaquatic Coleoptera collected in the Grand Portage Indian Reservation, Cook County, Minnesota, 2001 – 2011. BT = bottle trap, DN = dip net, LT = light trap, PFT = pitfall trap. Literature records: Haliplidae: Gundersen and Otrema, 1988; Dytiscidae: Gundersen and Christopherson 1990, Gundersen 1996, Larson et al. 2000, Hilsenhoff 1993; Gyrinidae: Ferkinhoff and Gundersen 1983, Majka and Kenner 2009; Hydrophilidae: Gundersen 2006, Smetana 1988; Elmidae: Hilsenhoff and Schmude 1992; Helodidae and Heteroceridae: McCafferty 1981; Chrysomeli- dae and Curculionidae: Merritt and Cummuns 1996. n.a. data not available.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes HALIPLIDAE (8) Haliplus blanchardi DN, June 2 (6), wide range of habitats Roberts Haliplus connexus DN, August, October 4 (3, 15), wide range of habitats Matheson Haliplus cribarius DN, June, July, August, 34 (3, 4, 7, 8, 10, 12, 15, 16), LeConte September widespread, shallow weedy habitats plus slowly flowing rivers Haliplus immaculicollis DN, May, June, July, 145 (1, 3, 4, 5, 6, 8, 10, 12, 16), Harris August, September common, wide range of habitats, except flowing water Haliplus leopardus DN, August 1 (3), wide spread, lentic habi- Roberts1 tats Haliplus longulus DN, LT, June 3 (7, 8), wide spread, common in LeConte shallow littoral habitats Peltodytes edentulus DN, June, September 2 (8, 16), wide spread, common (LeConte) in shallow littoral habitats Peltodytes tortulosus DN, June, July, August, 26 (2, 3, 4, 6, 8, 10, 15), north- Roberts September ern, shallow littoral habitats DYTISCIDAE (53) Acilius mediatus (Say) DN, July 1 (15), small forest pools with peaty substrate and adjacent stained creeks Acilius semisulcatus BT, DN, June, July, 20 (3, 7, 8, 10, 12), widespread, Aubé August, September many lentic habitats Acilius sylvanus BT, DN, June, July 2 (3), boggy lakes and ponds Hilsenhoff with emergent vegetation Agabus ambiguus (Say) DN, September 1 (17), springs, small streams and beaver ponds Agabus anthracinus BT, DN, May, June, 14 (2, 3, 6, 8, 9, 12), Mannerheim July, September widespread, common in permanent lentic habitats Agabus bifarius (Kirby) DN, May 1 (11), widespread, margins of vernal pools 2013 THE GREAT LAKES ENTOMOLOGIST 107

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Agabus confinis (Gyl- DN, September 3 (14), widespread, cool shaded lenhal) pools Agabus erichsoni Gem- DN, July 1 (16), widespread, dense veg- minger and Harold etation and margin of ponds Agabus phaeopterus DN, July 3 (17), margins of small pools (Kirby) with emergent vegetation Agabus semivittatus DN, June 1 (6), widespread, mostly lotic LeConte occasionally lentic Agabus seriatus (Say) DN, June, July, Sep- 14 (9, 14), widespread, springs tember and margins of small creeks Colymbetes sculptilis BT, DN, LT, June, July, 16 (3, 4, 6, 15 ), widespread, Harris August, September margins of lakes and ponds with emergent vegetation Coptotomus longulus BT, DN, LT, July, Au- 28 (3, 6, 10, 11, 12, 16), eastern, lenticus Hilsenhoff gust, September permanent ponds plus ditches and impoundments Desmopachria convexa LT, August 1 (n.a.), margins of bog and fen (Aubé) pools, also temporary ponds Dytiscus alaskanus J. BT, June, July 8 (3, 7), widespread, margins of Balfour-Browne permanent ponds and lakes Dytiscus cordieri Aubé BT, June, July 2 (10, 12), widespread, permanent, temporary and man-made habitats Dytiscus dauricus BT, DN, June 3 (6, 10, 11), widespread, perma- Gebler nent ponds including beaver ponds Dytiscus fasciventris BT, July, September 2 (10), ponds and marshes with Say extensive vegetation Dytiscus harrisi Kirby BT, July 1 (8), northern and eastern, large ponds and lakes Dytiscus verticalis Say BT, June, July 6 (2, 8, 11), northeastern, ponds, marshes and bogs Graphoderus fasciollis DN, June, July 3 (7, 10), northeast, permanent (Harris) ponds including beaver ponds Graphoderus liberus DN, June, August 9 (3, 10, 12), widespread, common (Say) in hardwood forests, rare in coniferous, boggy ponds and lakes Graphoderus perplexus BT, DN, June, July 21 (3, 6, 7, 8, 11, 12 ), wide- Sharp spread in boreal habitats, permanent ponds, lakes and bogs Hydaticus aruspex BT, DN, May, June 4 (3, 6, 7, 11, 12), widespread, Clark many lentic habitats 108 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Hydaticus piceus BT, DN, May, June, 4 (3, 12), permanent ponds, fens LeConte August and marshes Hydrocolus stagna- DN, June, July, Sep- 14 (6, 9, 11, 18), northeast, lis (Gemminger and tember margins of small cold pools and Harold) 1 creeks Hydroporus dentellus BT, LT, June, July, 13 (2, 6, 7, 8), widespread, bo- Fall August real, lentic habitats with dense vegetation Hydroporus dichrous LT, September 2 (11, 16), margins of small Melsheimer ponds and slow streams with dense vegetation Hydroporus fuscipennis BT, June 1 (8), widespread, small tempo- Schaum rary pools Hydroporus niger Say DN, July September 3 (n.a.), margins of sun-warmed pools Hydroporus notabilis DN, July 9 (6, 11), widespread, arctic and LeConte boreal, small pools Hydroporus tristis LT, August 5 (7, 8), small peatland pools Paykull and other lentic habitats Hygrotus falli (Wallis) 1 DN, June 2 (11), northeast and Great Lakes Hygrotus impresso- DN, June 1 (8), widely distributed, shal- punctatus (Schaller) low ponds and pools Hygrotus laccophilinus BT, LT, July, August 8 (3, 7), northeast, small ponds (LeConte) Hygrotus picatus BT, DN, May, June, 7 (3, 6, 7, 8), widespread, boreal, (Kirby) July ponds and marshes Hygrotus sayi DN, LT, May, June, 104 (3, 4, 8, 11), widespread, J. Balfour-Browne July, August, Septem- margins of ponds and slowly ber flowing streams Hygrotus tumidiventris BT, June 1 (11), temporary and perma- (Fall) nent ponds Hygrotus turbidus DN, June 2 (11), widespread (LeConte) Ilybius biguttulus BT, LT, June, August 2 (6), permanent warm lentic (Germar) habitats Ilybius fraterculus DN, June 1 (3), permanent ponds LeConte Ilybius picipes (Kirby) DN, June 1 (7), Permanent lentic habitats 2013 THE GREAT LAKES ENTOMOLOGIST 109

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Ilybius pleuriticus BT, DN, LT, June, July 9 (2, 4, 7, 8, 10), a wide range of LeConte lentic habitats Ilybius subaeneus DN, June, July 2 (2), shallow lentic habitats Erichson Lacccophilus biguttatus DN, May 1 (6), permanent ponds except Kirby peatland pools Lacccophilus maculosus BT, DN, June, July, 6 (8, 15), shallow permanent Say October pools and ponds Laccornis conoideus DN, October 1 (15), cold shallow pools, peat- (LeConte) land ponds and slow moving streams Neoporus undulatus BT, DN, June, July, 125 (3, 4, 6, 8, 10, 11, 16), (Say) August margins of beaver ponds, slow marshy streams and small lakes Neoscutopterus hornii BT, June, July 2 (1, 2, 3, 7), small mossy pools (Crotch) in bogs and fens Rhantus binotatus BT, DN, June, July, 5 (8, 14), primarily seeps, (Harris) September springs and small streams Rhantus consimilis DN, June, July 5 (11), warm weedy ponds Motschoulsky Rhantus sinuatus DN, September 1 (10), lentic habitats with (LeConte) dense emergent vegetation Rhantus wallasi Hatch DN, June 3 (3, 7), ponds with emergent vegetation GYRINIDAE (14) Dineutus assimilis DN, June, August 2 (3, 11), lakes and ponds Kirby Dineutus hornii Rob- DN, June, August, 80 (3, 6, 7, 16), lakes, ponds erts September and fens Dineutus nigrior DN, September 3 (6), bog lakes and other lentic Roberts habitats Gyrinus aeneolus DN, August, September 15 (4, 6, 7, 8, 14, 16), ponds and LeConte lentic lakes Gyrinus affinis Aubé DN, September 19 (14, 16), streams and lakes Gyrinus confinis DN, June, August, 96 (6, 7, 16, 19), mostly lentic, LeConte September also lotic habitats Gyrinus impressicollis DN, June, July 4 (6, 7, 9), lakes and streams Kirby Gyrinus latilimbus Fall DN, June, September 2 (8, 14), small lakes 110 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Gyrinus lecontei Fall DN, July, September 7 (6, 14), mainly small ponds and lakes Gyrinus maculeventris DN, June 1 (6), widespread, lentic habi- LeConte tats Gyrinus minutus Fa- DN, June, August 2 (6, 12), lentic habitats bricius Gyrinus pectoralis DN, June, July, August 65 (2, 3, 6, 11), lentic habitats LeConte Gyrinus pugionis Fall DN, June, August 4 (4, 16), mainly lentic habitats Gyrinus ventralis Kirby DN, June, September 3 (6, 8), mainly lentic habitats HYDROPHILIDAE (20) Anacaena lutescens DN, LT, July, August, 11 (4, 8, 14), ponds, marshes, (Stephens) September lake shores and stream margins Berosus seriatus (Say) DN, June 45 (4, 11), shallow lentic habitats or slowly moving water Berosus striatus (Say) DN, LT, May, June 33 (3, 4, 11), shallow lentic habitats or slowly moving water Cercyon roseni Knisch LT, June 2 (2, 3), margins of ponds, lakes, marshes and bogs Cymbiodyta vindicata DN, LT, June, August, 6 (7, 8), mainly lentic, stream Fall September edges, also some non aquatic habitats Enochrus hamiltoni DN, LT, May, June, 63 (1, 2, 4, 5, 7, 8), wide variety (Horn) July, August, Septem- of habitats, mainly shallow ber ponds with vegetation Enochrus ochraceus LT, August, September 6 (1, 2, 4, 5), ponds, lakes and (Melsheimer) small streams with dense vegetation Helophorus inflectus DN, May 5 (11), n. a. McCorkle Helophorus lineatus DN, July 1 (11), n. a. Say Helophorus sempervari- DN, September 2 (17), edges of shallow tempo- ans Angus rary pools and small streams Hydrobius fuscipes LT, PFT, July, August 3 (n. a.), bogs and shallow ponds (Linnaeus) Hydrobius melaenus LT, September 6 (1), mainly lotic habitats, oc- (Germar) casionally lentic Hydrochara obtusata BT, LT, June, August 12 (2, 4), shallow lotic and lentic (Say) habitats with dense vegetation 2013 THE GREAT LAKES ENTOMOLOGIST 111

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Hydrocus granulatus LT, June, July, Sep- 6 (8, 11), temporary shallow Blatchley tember ponds Hydrocus pseudosqua- DN, August 1 (10), n.a. mifer Mille Paracymus subcupreus DN, June, August 7 (8, 16), edges of ponds and (Say) lakes with emergent vegetation Tropisternus glaber DN, October 1 (14), marshes, swamps and (Herbst) bogs Tropisternus lateralis DN, LT, May, June 16 (2, 3, 4, 10, 16), lentic and nimbatus (Say) lotic habitats September Tropisternus mixtus DN, June, July, August, 48 (2, 3, 4, 6, 7, 8, 10, 12 LeConte September 14, 16), mostly ponds and shallow lakes, also stream margins Tropisternus natator (d’ DN, June, July 6 (2, 3, 4, 10), wide range of Orchymont) habitats, mostly ponds, also stream margins ELIMIDAE (3) Dubiraphia quadrino- DN, September 7 (17, 18), rocky riffles of small tata (Say) to medium-sized streams Optioservus fastitidus DN, June, September 4 (6, 18), rocky stream riffles (LeConte) and occasionally wave-swept lake margins Stenelmis crenata (Say) DN, September 1 (18), rocky stream riffles HETEROCERIDAE (1) Heterocerus brunneus LT, June 1 (4), margins of streams and Melsheimer rivers HELODIDAE (1) Cyphon nr. variabilis LT, August, September >500 (1, 2, 3, 5), larvae aquatic, Thurnberg adults terrestrial near water CHRYSOMELIDAE (7) Calligrapha philadel- DN, July 1 (17), reported on Cornus phica (L.) Donacia hirticollis DN, June, August 5 (3, 8), lentic habitats with Kirby vascular hydrophytes Donacia piscatrix DN, June 6 (3, 6, 7, 8), lentic habitats with Lacordaire vascular hydrophytes Donacia porosicollis DN, June, September 2 (2, 8), lentic habitats with Lacordaire vascular hydrophytes 112 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Table 1. Continued.

Family (no. sp.), Method and month(s) No. records (no. habitat species collected types), and literature notes Donacia tuberculifrons DN, June, July, August, 3 (6, 8, 16), lentic habitats with Schaeffer September vascular hydrophytes Neohaemonia nigricor- DN, June 14 (3, 4, 7, 8), lentic habitats nis (Kirby) with vascular hydrophytes Pyrrhalta decora (Say) DN, June, August 2 (3, 8), reported on Salix and Populus CURCULIONIDAE (5) Listronotus appendicu- DN, August 2 (16), lentic habitats with vas- latus (Boheman) cular hydrophytes Listronotus echinodori DN, June 2 (6, 11), lentic habitats with O’Brien vascular hydrophytes Listronotus grypidiodes DN, September 1 (8) lentic habitats with vascu- (Dietz) lar hydrophytes Lixus caudifer LeConte DN, July 2 (11), lentic habitats with vascular hydrophytes Notiodes limatulus DN, September 2 (11), lentic habitats with vas- (Gyllenhal) cular hydrophytes

1 new Minnesota state record. each. Based on these collections, seventy seven percent of species were considered rare or uncommon (1-10 records), twenty percent as common (11-100 records) and only three percent as abundant (more than 100 records). Sixty nine percent of species were collected at only 1 or 2 habitat types, twenty two percent at 3-5 and only nine percent at 6-10 habitats types (Table 1). However, caution must be used in interpreting both abundance and habitat data due to differences in sampling methods and unequal numbers of site collections.

Discussion HALIPLIDAE. Eight of the seventeen species of Haliplidae known to occur in Minnesota (Gundersen and Otremba 1988) were collected at the GPIR. Haliplus leopardus Roberts is recorded from Minnesota for the first time. The two most abundant species and habitat generalists, were H. immaculicollis Har- ris and H. cribarius LeConte. Haliplus immaculicolis occurs in a wide range of aquatic habitats and is widespread across Minnesota (Gundersen and Otremba 1988). Haliplus cribarius, found primarily in northeastern Minnesota, inhabits shallow weedy habitats. Three or fewer specimens of H. blanchardi Roberts, H. connexus Matheson and Peltodytes edentulus (LeConte) were collected at the GPIR. Gundersen and Otremba (1988) recorded H. connexus from north central and central Minnesota but not from the northeastern part of the state. Peltodytes edentulus, common and widespread across Minnesota, was rare at the GPIR. DYTISCIDAE. Sixteen genera and 53 species of Dytiscidae were recorded from the GPIR. In addition, one female specimen was tentatively identified as either Hydrocolus paugus (Fall) or H. persimilis (Crotch), another as either Hydroporus gossei Larson & Roughly or Hydroporus rectus (Fall) and two others 2013 THE GREAT LAKES ENTOMOLOGIST 113 as members of the Hydroporus obscurus Group, possibly H. obscurus (Sturm). Hydrocolus stagnalis (G. and H.) and Hygrotus falli (Wallis) were recorded for the first time from Minnesota. Based on distribution records for Minnesota Dytiscidae (Gundersen 1997a, 1997b; Larson et al. 2000), the following species are recorded for the first time from either the coniferous region or northeastern Minnesota: Acilius mediatus (Say), Dytiscus alaskanus J. Balfour-Browne, Hydaticus piceus LeConte and Neoscutopterus hornii (Crotch). Except for a pre-1940 record, this is only the second record of Graphoderus fasciollis from the coniferous region of Minnesota. Additional records for extreme northeastern Minnesota include Colymbetes sculptilis Harris, Desmopachria convexa (Aubé), Dytiscus cordieri Aubé, D. fasciventris Say, D. harrisi Kirby, D. verticalis Say, Graphoderus liberus (Say), Hygrotus impressopunctatus (Schaller), H. laccophilinus (LeConte), H. tumidiventris (Fall), Ilybius fraterculus LeConte, I. picpies (Kirby), Lacccophilus biguttatus Kirby, Laccornis conoideus (LeConte), Rhantus consimilis Motschoulsky and R. sinuatus (LeConte). The close proximity of the GPIR to Canada made it possible to evaluate broad habitat preferences of dytiscid species found at the GPIR based on their presence or absence in Canadian ecozones (Ecological Stratification Working Group 1996, Larson et al. 2000). Based on Larson et al. (2000, see Table 2) 28 species present at the GPIR occur in 9 or more ecozones while 14 species occur in 5 or fewer ecozones. Of those judged to be habitat specialists, Agabus semivittatus LeConte was the most specialized. GYRINIDAE. Dineutus hornii Roberts, Gyrinus affinis Aubé, G. confinis LeConte and G. pectoralis LeConte, the four most abundant species at the GPIR, are common and widespread across Minnesota and the upper Midwest (Ferkinhoff and Gundersen 1983). The few records of Dineutus assimilis Kirby and D. nigrior Roberts collected from the GPIR, most likely did not reflect their abundance as both are common in Minnesota but was likely due to the relatively few lotic habitats sampled. HYDROPHILIDAE. Eleven genera and 20 species of Hydrophilidae were recorded from the GPIR. While not included in Table 1, a larva of the genus Laccophilus was collected in 2003. Nearly 70% of the hydrophilid species collected at GPIR were rare to uncommon (< 10 records). Only Berosus striatus (Say), Enochrus hamiltoni (Horn) and Tropisternus mixtus LeConte were relatively common. Thirteen hydrophilid species were found at only 1 or 2 habitat types while two species were found at more than six different habitat types. Enochrus hamiltoni, a widely distributed and highly polymorphic species (Gundersen 1977, 2006), was collected at 10 GPIR sites. Cercyon roseni Knisch, Enochrus ochraceus (Melsheimer), Hydrocus granulatus Blatchley, Hydrobius fuscipes (Linnaeus) and H. melaenus (Germar) were collected only by light traps. C. roseni, a northern transcontinental species that is uncommon in Minnesota (Smetana 1978), was recorded from two sites. Cymbiodyta vindicata, uncommon in coniferous areas of the state (Gundersen 2006), was recorded from three sites. Both roseni and vindicata have been reported from semiaquatic as well as aquatic habitats (Smetana 1978, 1988). Hydrobius fuscipes occurs in shallow ponds and sphagnum bogs and H. melaenus in spring-fed streams and occasionally in standing water (Gundersen 2006). CHRYSOMELIDAE, ELIMIDAE, HETEROCERIDAE, HELODI- DAE and CURCULIONIDAE. Five genera and seven species of semiaquatic Chrysomelidae were collected at the GPIR, the most common of which was Neo- haemonia nigricornis (Kirby). The few riffle beetles collected at the GPIR were likely due to the relatively few lotic habitats sampled. Optioservus fastitidus, was collected from a wave-swept lake margin (Hilsenhoff and Schmude 1992) as well as streams and small rivers. The families Heteroceridae, and Helodidae were each represented by a single species. Cyphon variabilis (Helodidae), was common to very abundant in light trap collections. Adults of two non indigenous 114 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 root feeding weevils, Phyllobius oblongus (L.) and Polydrusus sericeus (Schaller), collected by dipnet samples in 2012, were not included as these species are not considered to be aquatic or semiaquatic (Bright 2006). However, the presence of oblongus and sericeus in dipnet samples suggests the possibility that these species utilize speckled alder [(Alnus incana (L.)] as a host plant (Pinski et. al 2005) that grows along swampy lake margins and wetlands.

Acknowledgments I wish to express my gratitude to Ralph Gundersen, Ethan Bright, and Paul Tinerella for their invaluable help in verifying the identity of some species of Coleoptera. I also wish to thank Margaret Watkins, water quality specialist, Andrew Schmidt, Wetland Specialist and Curtis Gagnon, Trust Lands Adminis- trator, of the Grand Portage Band for their help and permission to publish these results. I also thank my wife Bonnie and daughter Julie for their support and help.

Literature Cited Albert, D. A. 1994. Regional Landscape Ecosystems of Michigan, Minnesota, and Wis- consin: A working Map and Classification. U.S. Deptartment of Agriculture Forest Service General Technical Report. NC-178. 250 pp. Bright, E. 2006. Aquatic insects of Michigan. Aquatic Coleoptera (Beetles) of Michigan. Available from http://www.Insects.ummz.Isa.umich.edu/~ethanbr/aim/sp/Coleoptera/ sp (accessed 16 December 2010). Daussin, D. L. 1979. The aquatic coleoptera of Minnesota: a species list with references on taxonomy, biology, and ecology (Coleoptera: Haliplidae, Gyrinidae, Dytiscidae, Hydro- philidae, Hydraenidae, Elmidae, Dryopidae, Psephenidae, Helodidae, Chrysomelidae, Curculionidae). Newsletter Association of Minnesota Entomologists. 12(4): 6-18. Ecological Stratification Working Group. 1996. A National Ecological Framework for Canada. Agricultural and Agri-Food Canada, Research Branch, Centre for Land and Biological Resources Research and Environment Canada, State of the Environment Directorate, Ecozone Analysis Branch, Ottawa. 125 pp. Ferkinhoff, W. D., and R. W. Gundersen. 1983. A key to the whirligig beetles of Min- nesota and adjacent states and provinces (Coleoptera;Gyrinidae). Scientific Publica- tions of the Science Museum of Minnesota. New Series 5(3): 53 pp. Gundersen, R. 1977. New species and taxonomic changes in the genus Enochrus (Cole- optera): Hydrophilidae). The Coleopterists Bulletin 31 (3): 251-272. Gundersen, R. 1996. Dytiscidae of Minnesota I Dytiscinae, Cybistrinae. Occasional Papers in Aquatic Biology, St. Cloud State University 3(2): 40 pp. Gundersen, R. 1997a. Dytiscidae of Minnesota II Laccophilinae, and Colymbetinae, except Agabus. Occasional Papers in Aquatic Biology, St. Cloud State University 4(1): 54 pp. Gundersen, R. 1997b. Minnesota Dytiscidae III: Methlinae and Hydroporinae, exclusive of the Hydroporus group. Occasional Papers in Aquatic Bioliogy, St. Cloud State University 4(2): 45 pp. Gundersen, R. 2006. Hydrophilidae of Minnesota I Chaetarthriinae, Hydrobiinae, and Hydrophilinae (Coleoptera: Hydrophilidae). Occasional Papers in Aquatic Biology, St. Cloud State University 8(1): 78 pp. Gundersen, R. W., and C. Otremba. 1988. Haliplidae of Minnesota. Scientific Publica- tions of the Science Museum of Minnesota. New Series 6(3): 43 pp. Gundersen, R. W, and A. Christopherson. 1990. Dytiscidae of Douglas County. Oc- casional Papers in Aquatic Biology, St. Cloud State University. 1(1): 23 pp. 2013 THE GREAT LAKES ENTOMOLOGIST 115

Hilsenhoff, W. L. 1993. Dytiscidae and Noteridae of Wisconsin (Coleoptera). II. Distribu- tion, habitat, life cycle, and identification of species of Dytiscinae. The Great Lakes Entomologist 26(2): 35-53. Hilsenhoff, W. L. 1995. Aquatic insects of Wisconsin: keys to Wisconsin genera and notes on biology, habitat, distribution and species. Publication No. G3648. University of Wisconsin Natural History Museums Council. Madison, WI. 52 pp. Hilsenhoff, W. L., and K. L. Schmude. 1992. Riffle beetles of Wisconsin (Coleoptera: Dryopidae, Elmidae, Lutrochidae, Psephenidae) with notes on distribution, habitat, and identification. The Great Lakes Entomologist 25(3): 191-209. Larson, D. J. 1987. Aquatic Coleoptera of peatlands and marshes in Canada, pp. 99- 132. In D. M. Rosenberg and H. V. Danks (eds.), Aquatic Insects of Peatlands and Marshes in Canada. Memoirs of the Entomological Society of Canada No 140. 174 pp. Larson, D. L., Y. Alarie, and R. E. Roughley. 2000. Predaceous diving beetles (Co- leoptera: Dytiscidae) of the Nearctic Region with emphasis on the fauna of Canada and Alaska. NRC Research Press, Ottawa, Ontario, Canada. 982 pp. Majka C. G., and R. D. Kenner. 2009. The Gyrinidae (Coleoptera) of the Maritime Provinces of Canada: new records distribution, and faunal composition. ZooKeys 22: 355-372. Marschner, F. 1974. The original vegetation of Minnesota (map). Technical report, U. S. D. A. Forest Service, North Central Forest Experiment Station, St. Paul, MN. McCafferty, W. P. 1981. Aquatic Entomology The Fishermen’s and Ecologists’ Guide to Insects and Their Relatives. Jones and Bartlett. 448 pp. Meritt, R. W., and K. W. Cummins 1996. An Introduction to the Aquatic Insects of North America 3rd ed. Kendall/Hunt Pub. 862 pp. Pinski, R. A., W. J. Mattson, and K. F. Raffa. 2005. Host breadth and ovipositional behavior of adult Polydrusus sericeus and Phyllobius oblongus (Coleoptera: Curcu- lionidae), nonindigenous inhabitants of northern hardwood forests. Environmental Ecology 34(1): 148-157. Smetana, A. 1978. Revision of the subfamily Sphaeridiinae of America north of Mexico (Coleoptera: Hydrophilidae) Memoirs of the Entomological Society of Canada No. 105. 292 pp. Smetana, A. 1988. Review of the family Hydrophilidae of Canada and Alaska (Coleoptera) Memoirs of the Entomological Society of Canada No. 142. 316 pp. 116 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Epizooic Diatoms on the Cerci of Ephemeroptera (Caenidae) Naiads Daniel E. Wujek1

Abstract Using scanning electron microscopy, epizooic diatoms were observed growing on the cerci of Caenis amica Hagen naiads (Ephemeroptera, Caenidae). Meridion circulare (Greville) C. Agardh was the most abundant, followed by Synedra rumpens Kützing, then Cocconeis pediculus Ehrenberg. Other diatom species observed from substrates in Cedar Creek, Isabella County, Michigan were not observed on the cerci. No diatoms were observed on Ephemerellidae naiads.

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Epizoic algae are not infrequent on the surfaces of most active animals. The association between sloths and epizooic algae was one of the first to be described showing this unusual and interesting association since such relationships were described (Kuhn, in Welcher 1864). Freshwater epizootic algae have been described from a variety of hosts. These include protistans (Wiley et al. 1970, Pérez-Martinez et al. 2001); invertebrate animals such as Cladocera (Gaiser and Bachmann 1993, Barea-Arco et al. 2001), Rotifera (Wujek 2006); insects including Trichoptera (Bergey and Resh 1994, Sheath et al. 1995), Diptera (Sheath et al. 1996), and Odonata (Wujek personal observation); Crustacea together with Copepods (Russell and Norris 1971) and crayfish (Fuelling et al. 2010). The vertebrate hosts for epizootic algae include amphibians (Tumlison and Trauth 2006), reptiles including lizards and turtles (Gradstein and Equihua 1995, Garbary et al. 2007), fish (Shin et al. 2004), and captive polar bears (Lewin and Robinson 1979, Lewin et al. 1981).

Materials and Methods Spring collections containing naiad Ephemeroptera of the families Caenidae (N = 22) and Ephemerellidae (N = 43) were taken from a heavily shaded 200 m section of Cedar Creek, Isabella County, Michigan (34ºN33´794´´, 84ºW54´043´´) located between Wing and Deerfield Roads. The insects used for SEM observations, collected weekly from February through March 1988, were picked from substrates and immediately placed in buffered 7.2 pH 2% glutar- aldehyde. Insects used for light microscopy were fixed in 5% formalin. Insects which had been affixed to aluminum stubs were dehydrated, critical point dried, and sputter-coated with gold. Observations were done using an AMR 1200 scanning electron microscope. Diatom slides for light microscopic observations were prepared on both naiad families and material taken from various substrates associated with Cedar Creek where the mayflies were collected. These slides were observed using a Zeiss Photoscope II.

1Department of Biology, Central Michigan University, Mt. Pleasant, MI 48859. (e-mail: [email protected]). 2013 THE GREAT LAKES ENTOMOLOGIST 117 -

- (M), and and (M), (S). Scale Scale (S). naiad cerci cerci naiad naiad cercus cercus naiad A higher magnifica higher A Overview of diatoms diatoms of Overview Cocconeis pediculus Cocconeis . Caenis 1 Meridion circulare Meridion Caenis amica Caenis Figure on commu epizootic the showing µm. 20 = bar Scale nity. 2. Figure a of tion genera diatom epizooic with including (C), rumpens Synedra µm. 20 = bar 118 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Results and Discussion Three species of epizooic diatoms were observed growing abundantly only on the external surfaces of naiad mayfly cerci of Caenis amica Hagen, family Caenidae. They included Cocconeis pediculus Ehrenberg, Meridion circulare (Greville) C. Agardh, and Synedra rumpens Kützing (Figs. 1 and 2). By far the most numerous were cells of M. circulare. No diatoms were observed on the cerci of members of the family Ephemerellidae. The genus Meridion is instantly recognizable by most phycologists. It is an araphid, pinnate diatom found in both lentic and lotic habitats. Frustules are heteropolar in both valve and girdle views. In valve view, frustules appear clavate with a subcapitate footpole. Size ranges have been reported from 7-80 µm in length and 4-21 µm wide at the widest point. Costae are well developed. The species M. circulare has been described as a cosmopolitan taxon, although relatively rare in the tropics. It generally achieves maximal abundance in the spring, the season when the naiads for this study were collected. The species is considered to be alkaliphilous. It was also the most abundant species observed taken from natural substrates samples. Like M. circulare, Synedra rumpens is a cosmopolitan tychoplankton- benthic species. It is reported from many freshwater lakes or ponds or slow- moving streams (Patrick and Reimer 1986), most often from waters having a pH above 7.0. It was observed infrequently from the natural substrate samples. Cocconeis pediculus, like M. circulare, is very common and often is quite abundant in lowland streams and rivers. It also occurs in large numbers as an epiphyte in samples scraped from stones or woody debris. Its presence in such samples is probably related to the abundance of filamentous algae such as Cladophora at such sites. It favors circumneutral to alkaline water, often with high nutrient concentrations and also is able to tolerate moderate levels of organic pollution. A summary of its autecology can be found in Jahn et al. (2009). Other diatoms including Achnanthes lanceolata (Brébisson) Grunov A. minutissima Kützing, Fragilaria pinnata Ehr., Gomphonema olivaceum (Hor- nemann) Brébisson, Staurosirella leptostauron, var. dubia (Grun.) Edlund, and Planothidium hauckianum (Grun.) Round and Bakhiyarova were observed as periphyton or epilithon on substrates in the creek. They were not observed as part of the naiad’s epizooic flora. All are representatives of oligotrophic to me- sotrophic conditions and, as the three species observed on the cerci, considered alkaliphilous (Lowe 1974).

Acknowledgments Thanks to Tom Coppola for collecting the samples, Brian Roberts for assistance in the preparation of the illustrations, Drs. Rex Lowe and Mark Edlund for confirmation of some of the diatom identifications, and Dr. Donna King for identifying the naiad families and genus.

Literature Cited Barea-Arco, J., C. Pérez-Martinez, and R. M. Morales-Baquero. 2001. Evidence of a mutualistic relationship between an algal epibiont and its host, Daphnia pulicaria. Limnology and Oceanography 46: 871-881. Bergey, E. A., and V. H. Resh. 1994. Interactions between a stream caddisfly and the algae on its case: factors affecting algal quantity. Freshwater Biology 31: 153-163. Fuelling, L. J., J. A. Adams, R. J. Bixby, C. L. Caprette, H. E. Caprette, W. B. Chi- asson, C. L. Davies, M. M. Hall, W. L. Perry, M. L. Vis, and R. G. Verb. 2010. An occurrence of epizoic red algal chantransia on rusty crayfish (Orconnectes rusticus Giradr.) in west-central Ohio. Ohio Journal of Science 110: A-11. 2013 THE GREAT LAKES ENTOMOLOGIST 119

Gaiser, E. E., and R. W. Bachmann. 1993. The ecology and taxonomy of epizoic diatoms on Cladocera. Limnology and Oceanography 38: 6626-637. Garbary, D. J., G. Bourque, T. B. Herman, and J. A. McNeil. 2007. Epizoic algae from freshwater turtles in Nova Scotia. Journal of Freshwater Ecology 22: 677-685. Gradstein, S. R., and C. Equihua. 1995. An epizoic bryophyte and algae growing on the lizard Corythophanes cristatus in Mexican rain forest. Biotropica 27: 265-268. Jahn, R., K. Wolf-Henning, and O. E. Romero. 2009. Cocconeis pediculus Ehrenberg and C. placentula var. placentula (Bacillariophyceae): Typification and taxonomy. Fottea 9: 275-288. Lewin, R. A., and P. T. Robinson. 1979. The greening of polar bears in zoos. Nature 278: 445-447. Lewin, R. A., P. A. Farnsworth, and G. Yamanaka. 1981. The algae of polar bears. Phycologia 20: 303-314. Lowe, R. L. 1974. Environmental requirements and pollution tolerance of freshwater diatoms. Environmental Protection Agency, EPA-670/4-74-005. Patrick, R., and C. W. Reimer. 1966. The diatoms of the United States, exclusive of Alaska and Hawaii. Vol. 1. Academy of Natural Sciences of Philadelphia, Philadelphia. Pérez-Martinez, C., J. Barea-Arco, and P. Sánchez-Castillo. 2001. Dispersal and colonization of the epibiont alga Korshikoviella gracilipes (Chlorophyceae) on Daph- nia pulicaria (Cladocera). Journal of Phycology 37: 724-730. Russell, D. J., and R. E. Norris. 1971. Ecology and taxonomy of an epizoic diatom. Pacific Science 25: 357-367. Sheath, R. G., K. M. Müller, and D. J. Larson. 1995. Incorporation of freshwater Rhodophyta into the cases of caddisflies (Trichoptera) from North America. Journal of Phycology 31: 889-896. Sheath, R. G., K. M. Müller, M. H. Colbo, and K. M. Cole. 1996. Incorporation of freshwater Rhodophyta into the cases of chironomid larvae (Chironomidae, Diptera) from North America. Journal of Phycology 32: 949-952. Shin, J. K., H. S. Jeong, and S. J. Hwang. 2004. Ecological studies of epizoic algae attached on freshwater fishes in a small stream (Ian Stream), South Korea. Korean Journal of Limnology 37: 462-468. Tumlison, R., and S. E. Trauth 2006. A novel facultative mutualistic relationship between Bufonid tadpoles and flagellated green algae. Herpetological Conservation Biology 1: 51-55. Welcher, H. 1864. Ueber der entwicklung und den bau der haut und der haare bei Bradypus. Abhandlugen der Naturforschenden Gesellschatdten zu Hall. 9: 3-72. Wiley, R. L., W. R. Bowen, and E. Durban. 1970. Symbiosis between Euglena and damselfly nymphs is seasonal. Science 170: 80-81. Wujek, D. E. 2006. The first occurrence of the chrysophyte algaAmphirhiza epizootica from North America. Michigan Botanist 45: 197-200. 120 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Distribution records for Zenoa picea (Palisot de Beauvois, 1805) (Coleoptera: Callirhipidae) from the United States. Edwin L. Freese1

Abstract The known range of Zenoa picea (Palisot de Beauvois) (Coleoptera: Cal- lirhipidae) is summarized based on data from specimens, literature, and internet sources. This species, the only known member of the Callirhipidae occurring north of Mexico and previously thought uncommon, is recorded from 26 states and the District of Columbia in the United States. Its documented range is from Florida north to New Jersey, west to Iowa and Texas. No records are known from Mexico or Canada.

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Zenoa picea (Coleoptera: Callirhipidae) was described by Ambroise Ma- rie Francois Joseph Palisot, Baron de Beauvois (1752-1820), in 1805, after his return to Paris from expeditions in America and Africa (Palisot de Beauvois 1805, Chase 1925, Griffin 1932, Merrill 1937, Thomas 2009, Hájek 2011; Hájek, personal communication). The original generic assignment was Melasis, by French entomologist Guillame-Antoine Olivier, derived from the Greek for “black” which describes the insect as does the species name picea (Palisot de Beauvois 1805). In 1835, Thomas Say transferred the species to his new genus Zenoa, Greek for “a stoic” (Say 1835). Today Melasis (Olivier, 1790) is included in Eucnemidae (Eschscholtz 1829) the false click beetles (Muona 2002). Family status of this species has changed over the years with the most recent change being from Rhipiceridae (Latrielle 1834) to Callirhipidae (Emden 1924) (Crow- son 1950, 1955, 1971, 1973; Kasap and Crowson 1975; Forbes 1926; Lawrence and Newton 1982, 1995; Lawrence et al. 2000; Young 2002; Hunt et al. 2007; Hájek 2011). Palisot de Beauvois (1805) originally included this species in the Sternoxes (Buprestidae). This species is the only North American member of the Callirhipidae (Young 2002) and is seldom covered in field guides and textbooks. Adults are shining dark brown to black, 11-15 mm long, and found under logs and bark and prefer dry upland woods (Hájek 2011); they may be short-lived and noc- turnal being attracted to lights (Young 2002). Males have pectinate antennae and females have serrate antennae (Palisot de Beauvois 1805); males are more commonly collected at lights while females are more often collected on or reared from rotten wood (Hoffman et al. 2002). Larvae are found in rotten or dead wood (Lawrence et al. 2000) “usually in the presence of white-rot fungi.” (Young 2002). Palisot de Beauvois (1805) indicated the species lives on old trunks. The objective of this paper is to update the known range distribution of Zenoa picea in the United States and to document its distribution in Iowa. Young (2002) indicated its range as “southeastern United States, northwest to southern Illinois.” After discovering this species in Iowa, far from its known

133493 “S” Avenue, Adel, Iowa 50002. (e-mail: [email protected]). 2013 THE GREAT LAKES ENTOMOLOGIST 121 range, I conducted a literature survey for additional records and contacted cura- tors of entomological collections requesting specimen label data for this species. Field studies were made in Iowa to determine the range of the species in that state. The results of those data collection efforts are reported here. Data are reported herein from Iowa specimens, institutional and private collections, and literature searches.

Methods Adult specimens were collected in Iowa during the field seasons of 2003- 2012. Several methods were used including black light and florescent light in upland woodlands and inspecting logs and tree trunks with a flashlight at nighttime, along with searching logs and under bark during the daytime. Data also were obtained from on-line websites of several insect collections. The literature was searched for references on Palisot de Beauvois, state records, and information on species habits and range. Collections within Iowa were perused for specimens and out-of-state institutional collections were contacted to obtain label data. The following institutions, museums, collections, and individuals were que- ried for label data for this paper (abbreviations are those used in the text below). ABSC = Mark Deyrup, Archbold Biological Station, Venus, Florida AVEC = Arthur V. Evans, Smithsonian, Washington, D. C., personal collection CASC = Norman Penny, California Academy of Sciences, Golden Gate Park, San Francisco CISC = Cheryl Barr, Essig Museum of Entomology, University of California, Berkely CNCI = Serge Laplante, Canadian National Collection of Insects, Ottawa, Ontario DAVC = Doug A. Veal, Marion, Iowa, personal collection ELFC = Edwin L. Freese, Adel, Iowa, personal collection FMNH = James H. Boone, Field Museum of Natural History, Chicago FSCA = Paul Skelley, Florida State Collection of Arthropods, Gainesville INHS = Paul P. Tinerella, Illinois Natural History Survey, Champaign; James N. Zahniser, University of Illinois at Urbana/Champaign, Champaign ISIC = Iowa State University insect collection, Ames (Edwin L. Freese) JODC = James O. Durbin, Marion, Iowa, personal collection JPGC = Jeffrey P. Gruber, Madison, Wisconsin, personal collection MJPC = Matt J. Paulsen, Lincoln, Nebraska, personal collection MSUC = Gary Parsons, A. J. Cook Arthropod Research Collection, Michigan State University, East Lansing OSEC = Donald C. Arnold, K. C. Emerson Entomological Museum, Oklahoma State University, Stillwater OSUC = Creighton T. Freeman, Charles A. Triplehorn Insect Collection, Ohio State University, Columbus PKLC = Paul K. Lago, University of Mississippi, University, personal collection SEMC = Zachary Falin, Snow Entomological Museum, University of Kansas, Lawrence; Jennifer C. Thomas, Natural History Museum, University of Kansas, Lawrence TAMU = Edward G. Riley, Texas A and M University, College Station 122 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

UALR = Jeff K. Barnes, University of Arkansas, Little Rock UCFO = Shawn L. Kelly, Department of Biology, University of Central Florida, Orlando UMMZ = Mark F. O’Brien, University of Michigan Museum of Zoology, Ann Arbor UMRC = Kristin B. Simpson, Enns Entomological Museum, University of Mis- souri, Columbia UMSP = Ralph W. Holzenthal, University of Minnesota, St. Paul UNSM = Federico C. Ocampo and Matt J. Paulsen, University of Nebraska State Museum, Lincoln USNM = Richard L. Hoffman, Virginia Museum of Natural History, Martinsville, records from United States National Museum, The Smithsonian, provided via copy of his paper in Banisteria (Hoffman et al. 2002) VMNH = Richard L. Hoffman, Virginia Museum of Natural History, Martinsville WIRC = Steven J. Krauth, Wisconsin Insect Research Collection, University of Wisconsin, Madison

Results Zenoa picea specimen records by state as obtained from collections, published sources, and the internet; data as conveyed to me by a given collection manager-curator or as gleaned from the published literature; number of specimens per date and location in brackets are reported below. ALABAMA: NEW STATE RECORD. Tuscaloosa, 14 July 1950 [2], B. D, Valentine (FMNH); Walker Co., Jasper, 13 July 1980 [1], T. King, blacklight trap (FSCA); Mobile Co., Mobile, 11 June 2010, male [1], Robert Lord Zimlich, http://bugguide.net/node/view/409944; Jefferson Co., Vestavia, 10 August 1980 [1], T. King (CNCI); Montgomery, 7 July [1], Van Dyke Collection (CASC). ARKANSAS: NEW STATE RECORD. Ashley, Faulkner, Lee, Monroe, Pulaski, and Washington counties (UALR); Carroll Co., Eureka Springs, 30 July 1980 [1], J. R. Heitzman, blacklight (FSCA). DELAWARE: NEW STATE RECORD. Specimen in alcohol [1] (CUAMD 2007). FLORIDA: Alachua Co., Gainesville, June 2003 [1], J. M. Leavengood, Jr.; 7 June 1994 [1], E. H. Nearns; 12-16 June 1989 [1], P. J. Landolt; 28-30 June 1985 [1], 10 July 1980 [1], L. R. Davis, Jr.; 23 June 1959 [1], 16 July 1962 [1], J. W. Perry, light trap; June 1967 [1], September 1967 [1], September 1968 [1], L. A. Hetrick; 15 August 1962 [1], L. Rogers, light; 23 July 1963 [1], R. P. Esser, light; 24 June 1978 [1], R. A. Belmont, blacklight trap; Alachua Co., Gainesville, San Felasco Preserve, 14 June 1995 [1], E. H. Nearns; Alachua Co., Gainesville, Hogtown Creek, 19 June 1988 [2], P. Skelley, light trap; Alachua Co., Gainesville, Doyle Conner Building, 6-8 August 1973 [1], 23 May 1978 [1], 1-4 September 1972 [1], F. W. Mead, blacklight trap; Alachua Co., Gainesville, Belville Hts., 1 July 1980 [1], 2 July 1980 [2], L. A. Stange, blacklight trap; Alachua Co., Paynes Prairie, 16 July 1973 [1], D. P. Wojcik, blacklight trap; Clay Co., Goldhead SP, 8 June 1986 [1], L. Cicero; Dixie Co., 24 June 2001 [1], D. Almquist; uv light, salt marsh; Liberty Co., Torreya SP, 16 July 1987 [1], Matthews and Skelley, light; Palm Beach Co., Belle Glade, 2 October 1976 [1], R. A. Belmont; Putnam Co., Little Orange Lake, 3 June 1984 [1], K. W. Vick, blacklight trap; Putnam Co., Welaka Forest, 28-31 July 1986 [1], J. B. Heppner; Sebring, 18 May 1946 [1], H. Mason; Leon Co., Tallahassee, June 2006 [2], D. T. Alquist, uv light, backyard, mesic with hardwood trees (FSCA); Osceola Co., Walt Disney World, Sand Pine Plantation, 13 June 1996 [1], S. Fullerton, blacklight trap; Orange Co., Walt Disney World, Epcot Parking Site, Bayhead, 12 June 1996 [1], 1 July 1996 [1], 2013 THE GREAT LAKES ENTOMOLOGIST 123 Z. Prusak, and S. Fullerton, blacklight trap; Lake Co., Clermont, Arrowhead Estates, 2 July 2004 [1], S. L. Kelly and S. M. Fullerton, uv light, residential, red maple swamp (UCFO); Highlands Co., Highlands Hammock St. Pk., 15 June 1965 [1], blacklight trap, L. and C. W. O’Brien; 23-25 June [19]81 [2], uv light trap, S. Peck (CNCI); Florida (Downie and Arnett 1996); Florida (Staines 1983); Florida (Hájek 2011). GEORGIA: Dekalb Co., Stone Mtn. area, 3 July 1970 [1], J. E. Wappes (FSCA); Georgia, USNM (Hoffman et al. 2002); Georgia (Hájek 2011). ILLINOIS: Starved Rock, 10 August 1924 [1]; Piatt Co., 18 July 1971 [1], Reaves and Hollander; Union Co., Wolf Lake, 5 August 1963 [1], H. S. Dybas (FMNH); Stoney Brook, 12 July 1958 [2], Selander and Bouseman; Cornland, 10 July 1943 [1], Ross and Sanderson, reared decayed log; Mascoutah, 17 July 1906 [2], in woods; Homer Park, 11 July 1927 [1], T. H. F and R. D. G., at light; Putnam Co., 10 August 1932 [2], M. O. Glenn; Urbana, B’field Woods, 28 July 1945 [8], H. H. Ross; Bolter Coll. Univ. of Ill. [4] (INHS); Macon Co., NW side of Decatur, 12 July 1980 [1], 29 July 1981 [1], 6 July 1982 [1], 17 July 1983 [1], P. Skelley, light; Urbana, 22 July 1927 [1], A. E. Miller, from rotten log (FSCA); southern Illinois, Benj. D. Walsh, under bark, larva and adult (Osten Sacken 1862; Packard 1889); Vermillion Co., 4 July 1961 [1] L. D. Collom (CNCI); Il- linois, female [1], Van Dyke Collection (CASC); C. Ill. (ISIC); southern Illinois (Young 2002); Illinois, USNM (Hoffman et al. 2002); Illinois (Hájek 2011). INDIANA: Tippecanoe Co., 16 August 1972 [1], 17 July 1963 [1], 16 Au- gust 1967 [1]; Tippecanoe Co., Lafayette, 27 June 1990 [1]; Lafayette, 25 June 1991 [1]; Lawrence, 2 July 1934 [1], 20 July 1967 [1], N. M. Downie (FMNH); Tippecanoe Co., W. Lafayette, 7 July 1981 [1], M. and N. Deyrup (ABSC); W. Co., 16 August 1945 [1] (INHS); Vinccennes, 12 July 1931 [1] (OSUC); Posey Co., 9 July 1923 [1] (UMMZ); Monroe Co., Bloomington, 23-24 July 1984 [1], F. N. Young, BLT; Indianapolis, Camp Belzer, BSA, 5 July 1966 [1], 11 July 1966 [1], C. E. White; Warren Co., 16 August 1966 [1], C. D. Kerst (FSCA); Tippeca- noe Co., 16 July 2006 [1], Brad Barnd, “This specimen collected at night from trunk of mulberry tree that had a decent sap flow.”, http://bugguide.net/node/ view/96003; same location, collector, uv light, 9 July 2007 [1], http://bugguide. net/node/view/162491; Montgomery Co., Shades State Park, 22 July 2006 [1], Brad Barnd (Brad Barnd, personal collection and communication); Vigo, Putnam, and Posey Counties; “scarce” (Blatchley 1910); Indiana (Downie and Arnett 1996); Indiana (Hájek 2011). IOWA: Henry Co., “Phipceridae Zenoa picea beneath logs” [Spelling error as published.] (King 1914); Wickham (1911) did not list as known from Iowa; Johnson Co., Saylorville Lake, 16-31 July 1983 [2], R. H. Schieferstein (FSCA); Johnson Co., Saylorville Lake, 16-31 July 1983 [2], R. H. Schieferstein (TAMU); Louisa Co., 5 August no year given [1], G. Warren (FMNH); Hájek (2011) and previous investigators missed published record of King (1914). Fifty two adults were collected in Iowa during this study. Three specimens were collected in southeastern Iowa at Shimek State Forest, Van Buren Co., 2 August 2003 by Doug A. Veal using a combination UV-florescent light (DAVC, JPGC) and a fifth specimen was collected at the same location and lights 22 August 2004 (DAVC). A fourth specimen was collected in southwest Iowa at Waubonsie State Park, Fremont Co., 16 July 2004, using the same combination light, DAV (DAVC). A sixth specimen was collected in central Iowa at Hardin City Woodland State Preserve, Hardin Co., 16 July 2005, by Edwin L. Freese, using a black light (DAVC). More Iowa specimens were collected at Waubonsie State Park on 13 July 2009 [1] (ELFC), 24 July 2009 [7] (ELFC, DAVC), 2 July 2010 [1] (DAVC), 27 July 2010 [1] (ELFC), 5 August 2010 [1] (DAVC), 1 August 2011 [1] (DAVC), 9 June 2012 [2] (ELFC, DAVC), 12 July 2012 [2] (DAVC, ELFC), using the same combination light; 8 August 2009 [2] and 4 August 2010 [1] by James O. Durbin with black light (JODC); 14 July 2010 [2] (DAVC, ELFC) and 124 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

27 July 2010 [2] (ELFC) on firewood pile using flashlight. Further specimens were collected at Sharon Bluffs State Park, Appanoose Co., 22 July 2010 [7] (ELFC) and 26 July 2011 [4], black light, ELF (these four specimens forwarded to Jiří Hájek, The National Museum, Prague, Czech Republic, for DNA testing); 5 July 2012 [10] (DAVC, ELFC). The most recent new county record, Dallas Co., Raccoon River watershed, 18 June 2012 [1] combination light and 6 July 2012 [1] security light, wooded back yard (ELFC). KANSAS: Lawrence, Topeka, and Tonganoxie (Popenoe 1877); Jefferson Co., 18 July 2006 [1], Zack Falin, UV light; Crawford Co., Pittsburg, 14 July 2007 [1], Glenn A. Salsbury, UV light; Crawford Co., Frontencac, 21 July 2008 [1], Glenn A. Salsbury, UV light (SEMC); Douglas Co., F. H. Snow [2], determined by J. W. Angell “There were no dates on these specimens, but they were probably collected in the late 1800’s as Professor Snow was the founder and curator of the entomology collection from 1870 to 1901.” (Jennifer C. Thomas, personal communication); Douglas Co., Lawrence, 7 July 1925 [3], W. Benedict; Anderson Co., Garnett, 24 July 1956 [1], D. S. Lang (SEMC); Pottawatomie Co., 21 July 1955 [1] (FSCA); Douglas Co., 1923 [1], W. J. Brown (CNCI); Johnson Co., Mission, 10 July 2009, male [1], porch light, 1950s suburb with assorted mature trees, Betsy Betros, http://bugguide.net/node/view/494970 (Betsy Betros, personal communication); Kansas (Staines 1983); Kansas (Hájek 2011). KENTUCKY: Hickman, July 1937 [1], E. E. Byrd (FMNH); specimen labeled only Kentucky [1] (UMMZ); “Murphy’s Pond, Hickman Co., KY., 1964, J. M. Campbell, Reared from larva coll. in oak stump; larva coll. IV-17-1964, Adults emerged VI-22-1964” [5] (CNCI); Kentucky, USNM (Hoffman et al. 2002); Kentucky (Hájek 2011). LOUISIANA: Jefferson Parrish., Kenner, 9 July 1956 [1], H. L. Dozier, lights; Baton Rouge, 24 June 1961 [1], Gary N. Ross; Edgard, 14 June 1973 [1], 29 June 1973 [1], 5 July 1973 [1], V. Brou, uv light; Sunshine, June 1972 [1], 5 June 1972 [1], 11 June 1972 [1], 13 June 1972 [2], 22 June 1972 [1], 3 August 1972 [1], 8 August 1972 [1], 9 August 1972 [1], 11 august 1972 [2], V. A. Brou, blacklight trap (FSCA); Assumption, Caddo, East Baton Rouge, Franklin, Iber- ville, Saint James, Saint John the Baptist, Saint Martin, Tensas, West Baton Rouge, and West Feliciana (LSAM 2009); East Baton Rouge Parish (TAMU); Madison Co., Mound [1] (UMSP); Lake Pontchartrain region (Sommers 1874); Norco, Bonnet Carre Spillway, St. Charles Parish, 30 May 1971 [2], 12 June 1971 [1], Eric H. Metzler, determined D. K. Young (MSUC); Madisonville, 1 July [19]83 [1], BF and JL Carr (CNCI); Caddo Parish, Shreveport, July 1973 [1], Cheryl B. Barr; Feliciana Parish, Tunica Hills, Weyanoke, 15 August 1986 [1], 26 June 1987 [1], C. B. Barr, BL and MV; St. Tammany Parish, Pearl River, 17 June 1982 [1], C. B. Barr and E. G. Riley, black light (CISC); Louisiana, USNM (Hoffman et al. 2002); Louisiana (Hájek 2011). MARYLAND: Anne Arundel Co., Edgwater, 27-29 July 1983 [1], 29 July-1 August 1983 [1], 30 July-11 August 1988 [1], 5-10 August 1988 [1], 13-16 July 1990 [1], 19-22 July 1991 [1], 16-18 July 1992 [2], 7-10 August 1992 [1], 20-27 July 1993 [1], 18-20 July 1994 [2], 20-22 July 1994 [1], 22-25 July 1994 [1], 25 July-1 August 1994 [1], C. L. Staines, Jr., blacklight (FSCA); Anne Arundel Co., Annapolis, 1 August 1988, male, [1], Frank Guarnieri, collected at light, http://bugguide.net/node/view/361079; Anne Arundel Co., 24 July 1995 [1], M. E. Epstein and WES, “At black light; open sandy gap in mixed forest”, USNM; Baltimore Co., Essex, 26 April 1987 [1], WES and J. M. Swearingen, under driftwood and debris on sand beach”, USNM; Charles Co., Smallwood State Park, 27 July 2002 [2], WES, J. M. Swearingen et al., “At black light in mature mixed forest”, USNM; Montgomery Co., Ashton, 27 July [1] and 30 July 2002 [1], G. F. Hevel; Cabin John Bridge, 28 July 1912 [1], F. Knab, USNM; Jacksons Island, 8 August 1920 [1], H. S. Barber, USNM; North Potomac, 6 July 2002 [1], Steven Lingafelter, blacklight, USNM; Plummer’s Island, 5 November 1905 [3], H. S. 2013 THE GREAT LAKES ENTOMOLOGIST 125 Barber, larvae, “in log of sassafras”, USNM; 22 July 1995 [2], M. E. Epstein, WES, and J. M. Swearingen, “In mixed forest; at black light on island summit”. USNM; 16 July 1997 [1], M. E. Epstein, WES, J. M. Swearingen, USNM; 13 July 2002 [2], WES, J. M. Swearingen et al., “On rotten wood at base of dead standing Quercus rubra at dark”, USNM; Rockville, 2 July 1980 [1], 26 July 1980 [1], Scott W. Gross, USNM; Prince Georges Co., Cedarville State Forest, 20 July 2002 [1], WES, J. M. Swearingen et al., “At black light at edge of mature mixed forest near open pond”, USNM; same location, date [1], collectors, “on rotten wood in hollow base of beech”, USNM; College Park, 18 July 1942 [1], George B. Vogt, light, USNM (Hoffman et al. 2002); Anne Arundel Co., Edgewater, 15 July 1980; Baltimore Co., Hebbville, 17 July 1969, Ten Hills, 22 July 1982, at light, R. S. Bryant collection; Harford Co., Aberdeen, 10 July 1963; Montgomery Co., Cabin John, 28 July 1912; Plummer’s Island, 19 August 1903, 15 August 1916, 6 August 1918 (Staines 1983); Maryland (Hájek 2011). MICHIGAN: NEW STATE RECORD. Wayne Co., Grosse Ile, 21 July 1956 [1], Geo. Steyskal (MSUC); No specimens in UMMZ collection (Mark F. O’Brien, personal communication). MISSISSIPPI: NEW STATE RECORD. Lafayette Co., Oxford, 2 August 1978 [1], 29 July 1988 [1], 26 June 1988 [1]; Issaquena Co., 2 mi SW Shipland, 20 June 1992 [2], Paul K. Lago (PKLC). MISSOURI: St. Louis, 24 June 1896 [1]; Columbia, 16 July 1904 [1]; Colum- bia, 16 July 1956 [1], M. S. Enns; Columbia, 6 July 1968 [4], 9 July [1] 1968, reared indoors from larvae from rotten log collected in April, W. S. Craig; Columbia, 17 July 1966 [1], P. L. Andrews; Columbia, 20 July 1948 [1], M. A. Rowoth; Colum- bia, 1 August 1968 [1], C. Garry; Boone Co., Columbia, 10 July 1967 [2], 11 July 1967 [1], 19 July 1967 [1], 20 July 1967 [1], 23 July 1967 [2], 25 July 1967 [1], 27 July 1967 [3], 29 July 1967 [1], 31 July 1967 [1], 6 August 1967 [4], light trap collecting; Boone Co., Columbia, 27 July 1976 [1], black light trap, 17 July 1977 [1], black light, S. E. Thewke; Boone Co., Columbia, 27 July 1978 [1], Marlin E. Rice; Boone Co., 29 June 1975 [2], south of Columbia, S. E. Thewke, blacklight; Boone Co., 1 August 1972 [1], S. O. Swadener; Boone Co., Ashland, 22 July 1968 [1], 4 August 1968 [1]; Boone Co., Ashland Wildlife Area, 9 August 1968 [1], light trap; 20 July 1978 [1], 5 August 1978 [1], G. Ulmer; Randolph Co., 1 mi. E. Moberly, 16 July 1972 [2], 29 July 1972 [1], T. G. Riley; 2 July 1977 [1], E. G. Riley; Howard Co., Bell’s Orchard, Boonville, 21 July 1957 [1], F. Wood; Cole Co., 18 July 1970 [1], Ramon Gass; Douglas Co., Vanzant, 19 July 1975 [1], 1 July 1977 [1]; Maries Co., 5 mi. NW Vienna, 12 May 1972 [1], S. E. Thewke, reared in lab; Jackson Co., 9 July 1967 [1], Leg. Roger L. Heitzman, UV; Kan. C. [1] (UMRC); St. Louis, 4 July 1907 [1], E. Liljeblad (UMMZ); Randolf Co. (TAMU); Jackson Co., July 1955 [1], Ronald H. Pine, determined by J. F. Lawrence (SEMC); St. Louis, 4 July 1907 [1], G. A. Akerlind; Stanton, 25 July 1966 [1], K. Stephan; Benton Co., 3 mi. NW of Warsaw, 3 July 1975 [1], R. L. Heitzman, uv light; Clay Co., Coolie Lake, 18 July 1968 [3], 17 May 1975 [1], 1 September 1986 [1], R. Heitzman, uv light; 17 June 1971 [4], G. Nelson and R. Heitzman, light; Jackson Co., Blue Springs, 16 July 1997 [1], G. H. Nelson, uv light; Jackson Co., Raytown, 5 July1973 [1], 7 July 1973 [1], 9 July 1973 [2], 11 July 1973 [3], 13 July 1973 [2], 16 July 1973 [2], 17 July 1973 [5], 18 July 1973 [5], 19 July 1973 [1], 20 July 1973 [1], 26 July 1973 [1], 27 July 1973 [6], 28 July 1973 [3], 29 July 1973 [2], G. H. Nelson, uv light; Morgan Co., Lake o’t Ozarks, 11 June 1982 [1], R. Heitzman, blacklight; Wayne Co., Markham Spring NF, 28 July 1986 [1], J. R. Heitzman; cane, cypress, tupelo, pine and deciduous forest (FSCA); Franklin Co., 1 July 1967 [1] (OSUC); Benton Co., Zenoa picea (Shockley and Cline 2004); Columbia, 15 July 1966 [1], Poe, at light (WIRC); One specimen badly damaged labeled only MO (MSUC); Stanton, 25 July [19]66 [1], K. Stephan (CNCI); Boone Co., west of Columbia, 26 June 2012, male [1], specimen not kept, black light, back yard with second growth oak-hickory woodland, Lee Elliott, http://bugguide.net/node/view/670041 (Lee Elliott, personal communication); Missouri, USNM (Hoffman et al. 2002); Missouri (Hájek 2011). 126 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

NEBRASKA: NEW STATE RECORD. Richardson Co., Indian Cave State Park Trail, 1 August 2003, male [1], at night in rotting stump, M. J. Paulsen; Nemaha Co. Indian Cave Trail, 14 July 2006, female [1], MV light, S. Spomer; Otoe Co., Lewis and Clark Visitor Center, 30 July 2007, male [2], lights, M. J. Paulsen (UNSM); Nemaha Co., Indian Cave Park, 17 July 2006 [1], UV light, MJP; Richardson Co., Indian Cave Trail, 17 June 2004, male [1], blacklight, M. J. Paulsen (MJPC); Sarpy Co., Fontenelle Forest, Bellevue, 9 July 2009, male [1], came to light, no specimen kept, Loren and Babs Padelford, http://bugguide. net/node/view/490001 (Babs Padelford, personal communication). NEW JERSEY: “Coleoptera: Mr. Kaeber exhibited ….. Zenoa picea Beauv. Collected at Red Banks, New Jersey, July 4, 1908.” (Greene 1915); “Zenoa picea Beauv. Red Bank, July 4, 1908. Kaeber.” (Weiss 1916); not listed by Hájek (2011). NORTH CAROLINA: 7 pinned specimens (NCSU 2011); Columbus Co., Lake Waccamaw, 6 July 1985 [1], WES and A. Gerberich “At black light in oak and pine scrub sand barrens near lake”, USNM (Hoffman et al. 2002); no known specimens (Brimley 1938); North Carolina (Hájek 2011). OHIO: Clinton Co., 18 July 1955 [1], 8 July 1961 [1], 9 June 1968 [1]; Richland Co., 20 July 1967 [1] (OSUC); Cincinnati, 12 July 1947 [1], D. H. Kistner; Columbus, 18 July 1962 [1], DJ and KN Knull; Champaign Co., 24 July 1954 [1]; Athens Co., 1 July 1956 [1], 19 July [1] 1956, L. W. Ressegger (FMNH); Hamilton Co., Cincinnati, 4 July 2012 [1], Steve Pelikan, http://bugguide.net/node/view/668860; Knox Co., 11 July 1941 [1], 16 July 1942 [1], H. F. Strohecker (FSCA); near Cincinnati (Dury 1902); Ohio (Downie and Arnett 1996); Ohio (Hájek 2011). OKLAHOMA: Blaine [1], Caddo [1], Carter [1], LeFlore [1], Noble [1], Osage [1], Pawnee [1], Payne [13], Sequoyah [1], Tulsa [1], and Woodward [1] counties, Dates range from June 14 to Aug. 13. Most were taken in black light traps.” (OSEC); Grady Co. (UALR); Latimer Co. (TAMU); Sequoyah Co., Vian, 14 May 1969 [1], S. Weaver, blacklight; Red Rock Canyon SP, 26 July 1975 [1], K. Stephan (FSCA); Summerfield, 14 June 1939 [1], Kaiser-Nailon (CNCI); Sequoyah Co., 1 July 1929 [2], R. D. Bird, R. Hopping Collection (CASC); Cleve- land Co., Norman, 3 July 1975 [1], 31 July 1975 [1], 8 August 1975 [1], 12 July 1976 [1], 18 July [1] 1976, William D. Shepard, leg, light trap; Okfuskee Co., 5 mi N Beardon, 25 July 1976 [2], William D. Shepard, leg, in rotten log (CISC); Oklahoma, USNM (Hoffman et al. 2002); Oklahoma (Hájek 2011). PENNSYLVANIA: Hummelstown, 14 July 1917 [1], J. N. Knull (FMNH); Phila, P. A., no date [1] (OSUC); Delaware Co. (TAMU); Cowens Gap SP, 10-11 July 1988 [1], E. Giesbert (FSCA); southwestern Pennsylvania, rare (Hamilton 1895); “Pittsburg[h], IX. 19 20 Pa” [1] (CNCI); Pennsylvania [1], Carl Fuchs Col- lection (CASC); Pennsylvania (Downie and Arnett 1996); Pennsylvania (Staines 1983); Pennsylvania (Hájek 2011). SOUTH CAROLINA: Florence Co. [1]; Pickens Co. [1]; Pickens Co., one in alcohol (CUAMD 2007); Florence Co., June-July, at lights, tanglefoot screen, under dead oak bark (Kirk 1969); Horry Co., Myrtle Beach, 22 August 1947 [1], T. H. Hubbell (FSCA); South Carolina (Hájek 2011). TENNESSEE: NEW STATE RECORD. Deer Lodge, 20 August 1932 [1], B. Benesch (FMNH); Deer Lodge, 25 June 1916 [1], C. Kreiger (UMMZ). TEXAS: Anderson, Bell, Brazoria, Brazos, Fort Bend, Kaufman, Lime- stone, Montgomery, Sabine, and San Jacinto counties (TAMU); Fort Bend Co. (UALR); Nacogdoches Co., Nacogdoches, 3 July 1994 [1], C. Ely (SEMC); Austin Co., Austin SP, 8-9 July 1980 [1], R. H. Arnett, Jr., blacklight trap, oak; Houston, 5 July 1980 [1], O. E. Hunt, light; Anderson Co., Engling WMA, 29 May 1998 [1], Godwin and Wappes, MV/UVL; Montgomery Co., Woodlands, 14-20 June 1977 [1], 25 June 1977 [1], 24-30 June 1979 [1], J. E. Wappes (FSCA); Waco, July 1968 [1], D. Kavanaugh (CNCI); Anderson Co., Blackfoot, Engeling Wildlife 2013 THE GREAT LAKES ENTOMOLOGIST 127 Management Area, 12 June 1994, male [1], Mike Quinn, http://bugguide.net/ node/view/453068; Texas, Victoria, “in morschen Stammen”, 20 December 1902 – 22 February 1910, 5 larvae, J. D. Mitchell, leg. (Emden 1932; Mark Deyrup, personal communication); Texas (Staines 1983); Texas (Hájek 2011). VIRGINIA: Middleburg, 7 July 1948 [1], H. L. Dozier (FSCA); Charles City Co., Harrison Lake National Fish Hatchery, 12 July 2006, male [1], C. E. Hatter; Prince William Co., Bull Run Mountains, 7 July 2006, male [1], 4 August 2006, female [1], uv trap, K. H. Bass and M. J. Kieffer; 29 July 2005, female [1], uv, M. J. Kieffer, K. H. Bass, and J. T. Warden; Surry Co., Chip- pokes State Park, 15 August 2007 [1], uv trap, basic mesic forest, A. C. Chazal and M. E. Dougherty (AVEC); Middlesex Co., 18 July 2000 [1], Malaise trap; Dickenson Co. [7]; City of Chesapeake [4] and City of Virginia Beach [1], “Robert Vigneault almost certainly by pulling off loose bark” (VMNH, Richard Hoffman, personal communication); Fairfax Co., Black Pond, 9-19 July no year given [1], F. C. Craighead, reared from Castanea dentata, USNM; Falls Church, 3 June 1917 [1], G. M. Greene, USNM; near Plummer’s Island, 27 July 1920 [1], H. S. Barber, USNM; King George Co., Chotank Creek Natural Area Preserve, Berthaville, 21 August 2001 [1], K. L. Derge and R. O. Wilson, uv light, VMNH; King and Queen Co., Dragon Run Swamp, 18 July 2000 [1], C. S. Hobson and A. C. Chazal, VMNH; Loudoun Co., Bluemont, 15 July 1911 [1], W. R. Walton, USNM (Hoffman et al. 2002); Virginia (Hájek 2011). WEST VIRGINIA: Morgan Co., 7 km east of Paw Paw, 25 July 1999 [1], Frank Guarnieri, uv and mercury-vapor light, http://bugguide.net/node/ view/361077; Hampshire Co., 10 km S. Capon Bridge at Cacapon River, 21-22 July 2002 [1], WES, J. M. Swearingen, J. R. Ott and E. Silverfine, “At black light in mixed mature forest, crest of shale slope above river”, USNM (Hoffman et al. 2002); West Virginia (Hájek 2011). DISTRICT of COLUMBIA: #1351 Zenoa picea Beauv. (Ulke 1903); Oxon Run, 16 July 1921 [1], E. V. Shannon, “on oak bark”, USNM (Hoffman et al. 2002); District of Columbia (Hájek 2011).

Discussion More than 475 specimen records were collected for Zenoa picea during this study from 26 states and the District of Columbia. Iowa, Illinois, Indiana, Ohio, Pennsylvania, and New Jersey now mark the northern edge of the known range and Nebraska, Kansas, Oklahoma, and Texas the western edge. No specimens were located from Minnesota, Wisconsin, South Dakota, and Canada (Ralph Holzenthal, Steve Krauth, Jeff Gruber, Rita Velez, Francois Genier, Serge Laplante, personal communication). Records from Alabama, Arkansas, Delaware, Michigan, Mississippi, Nebraska, and Tennessee represent new state records having not been cited by Hájek (2011); Hájek (2011) did not list Iowa as he and other investigators missed the record published by King (1914). The single Michigan specimen from 1956 was collected just south of Detroit near the Detroit River and thus across from Ontario, Canada, indicating the possibility that the species may occur in southern Canada. It may also be present in Mexico just south of Texas but no attempt was made to locate specimens south of the United States. Most specimens were collected south of 42 degrees north latitude and east of 99 degrees west longitude (Fig. 1). Month of collection was obtained from 390 specimens with adults being active from April through October. The earliest adult collected was 26 April [1987] in Baltimore Co., Essex, Maryland and the latest collected specimen was 2 October [1976] from Palm Beach Co., Belle Glade, Florida. Temporal distribution of adult collection date records were: April (1), May (7), June (60), July (266), August (51), September (4), and October (1). 128 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Figure 1. Range map of Zenoa picea.

Label data obtained indicate a few specimens were reported as having been reared from wood or found on rotten wood, a tree, stump, under bark, or under driftwood; most specimens were collected at lights or in a light trap. Many specimens were noted as collected in association with older trees in state forests, parks, and wildlife areas as well as national parks with label data of woodland, forest, mixed forest, basic mesic forest, mature mixed forest, and mixed mature forest. According to Hoffman et al. (2002) “the species is perhaps an indicator of mature forest habitat”. Prior to Euro-American settlement the Iowa collecting sites were probably dominated by grasslands and savanna on the uplands and forest habitat only along river corridors; today the sites are dense oak woodlands and forest (Smith 1998; Mutel 2008). The collection site at Shimek State Forest is an upland area about two miles from the Des Moines River and three miles from the Missouri state line; the surrounding forest contains mostly white, red, and black oaks (Mark Leoschke, personal communication); the State of Iowa started to purchase these lands in the 1930s. The dominant upland tree species at Waubonsie State Park is chinquapin oak (John Pearson, personal communication) with bur and white oak, basswood, paw paw, and many other tree species present. Pre-settlement vegetation of these loess hills was probably grassland and savanna (John Pearson, personal communication; Mutel 1989). The collection site is approximately five miles from the Missouri River and 5 miles from the Missouri state line. The State of Iowa purchased lands and founded this park in 1926 (Wolf 1991). Hardin City Woodland State Preserve contains forest on level upland, north-facing slope, and floodplain adjacent the Iowa River in 2013 THE GREAT LAKES ENTOMOLOGIST 129 central Iowa with red and black oak, shagbark hickory, black and sugar maple, and basswood being common (Freese 2005). The Iowa River at this location meanders through a deep U-shaped wooded valley of glacial origin (Freese 2005); the acreage for this preserve was purchased during 1959 (Herzberg and Pearson 1991). Sharon Bluffs State Park located along the Chariton River about nine miles from the Missouri border, contains a forest of white and pin oak and shagbark and kingnut hickory (Mark Leoschke, personal communication) with buckeye, basswood, and silver and sugar maple also present; the original first acreage for this park was purchased during 1929 (Wolf 1991). After collecting the first specimens in Iowa, far from the published known range (Young 2002), my first thought was that the species may now be spreading north and westward following human disturbance, with the beetle possibly spreading by migrating up the major river valleys (Mutel 2008; Smith 1998; Jungst et al. 1998; Schwert 1996; Cruden and Gode 2000; Schlicht et al. 2007; Turnbull 1980). But specimen data from many decades ago, especially from Kansas before 1877 (Popenoe 1877) and Iowa (King 1914), indicate the species may already have been this far west and north but just seldom collected and these western records were unknown or missed by researchers. Recent collecting throughout its range indicates Z. picea is present in forested areas across a large geographic area of the eastern and central United States but overall probably not in large numbers.

Acknowledgments First a special thank you to my collecting buddy, Doug Veal, for sharing his interest in beetles and being a great friend. The author also thanks all the cura- tors, collection managers, and collectors listed above who contributed data for this project. Greg Courtney kindly provided access to the Iowa State University insect collection. The author also thanks the following who contributed information to this study: Dan Young, Matt Paulsen, Paul Lago, Jiří Hájek, Warren Steiner, Mark Deyrup, John Pearson, Mark Leoschke, Eric Day, Steve Roble, Francois Genier, Becky Heth, Loren and Babs Padelford, Betsy Betros, Brad Barnd, Deb Lewis, Rita Velez, Lee Elliott, Kevin Thorne, the late Richard L. Hoffman, and all the contributors and creators at BugGuide.Net and Biodiversity Heritage Library. Also, a thank you to the four reviewers, their constructive comments have made this a better paper. And finally, a special thank you to my wife Janet, for her interest in nature, and thus understanding my interest in beetles.

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Crowson, R. A. 1971. Observations on the superfamily Dascilloidea (Coleoptera: Polyphaga), with the inclusion of Karumiidae and Rhipiceridae. Zoological Journal of the Linnean Society 50(1): 11-19. Crowson, R. A. 1973. On a new superfamily Artematopoidea of polyphagan beetles, with the definition of two new fossil genera from the Baltic Amber. Journal of Natural History 7: 225-238. Cruden, R. W., and O. J. Gode. 2000. The Odonata of Iowa. Bulletin of American Odonatology 6(2): 13-48. Downie, N. M., and R. H. Arnett. 1996. The Beetles of Northeastern North America. 2 Volumes. Sandhill Crane Press, Gainesville, Florida, 1,721 pp. Dury, C. 1902. A revised list of the Coleoptera observed near Cincinnati, Ohio, with notes on localities, bibliographical references, and descriptions of new species. The Journal of the Cincinnati Society of Natural History 20(3): 107-196. Available from http://www.biodiversitylibrary.org/item/44750 (accessed July 2011). Emden, F. van. 1924. Zur Kenntnis der Sandalidae. II. u. III. Entomologische Blät- ter 20: 86-99. Emden, F. van. 1932. Die Larven der Callirhipini, eine mutmassliche Cerophytum-Larve und Familien-Bestimmungstabelle der Larven der Malacodermata-Sternoxia-Reihe (Coleoptera) (Zur Kenntnis der Sandalidae 18). Bulletin et Annales de la Société Entomologique de Belgique 72: 199-259 + pls. XI-XIII. Forbes, W. T. M. 1926. The wing folding patterns of the Coleoptera. Journal of the New York Entomological Society 34(1): 42-68 and 34(2): 91-139. Freese, E. L. 2005. Vascular plant species list, Hardin City Woodland State Preserve, Hardin County, Iowa. Unpublished report to Iowa Department of Natural Resources, State Preserves Board, 16 pp. Greene, G. M. 1915. Feldman collecting social. Entomological News and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 26(1): 237-238. Available from http://www.biodiversitylibrary.org/item/84799 (accessed July 2011). Griffin, F. J. 1932. On the dates of publication of ‘Palisot de Beauvois, Insectes rec. Afr. Amer., 1805-(1821). Annals and Magazine of Natural History 10: 585-586. Hájek, J. 2011. World catalogue of the family Callirhipidae (Coleoptera: Elateriformia), with nomenclatural notes. Zootaxa 2914: 1-66. Hamilton, J. 1895. Catalogue of the Coleoptera of Southwestern Pennsylvania, with notes and descriptions. Transactions of the American Entomological Society 22: 317-381. Available from http://www.biodiversitylibrary.org/item/32378 (accessed July 2011). Herzberg, R., and J. A. Pearson. 2001. The Guide to Iowa’s State Preserves. Univer- sity of Iowa Press, Iowa City, 214 pp. Hoffman, R. L., S. M. Roble, and W. E. Steiner, Jr. 2002. Thirteen additions to the known beetle fauna of Virginia (Coleoptera: Scirtidae, Bothrideridae, Cleridae, Te- nebrionidae, Melyridae, Callirhipidae, Cerambycidae, Chrysomelidae). Banisteria 20: 53-61. Available from http://www.cmi.vt.edu/vnhs/banisteria/pdf-files/ban20/ Banisteria%2020New%20Beetles.pdf (accessed March 2011). Hunt, T., J. Bergsten, A. Levkanicova, A. Papadopoulou, O. St. John, R. Wild, P. M. Hammond, D. Ahrens, M. Balke, B. Michael, M. S. Caterino, J. Gomez- Zurita, I. Ribera, T. G. Barraclough, M. Bocakova, L. Bocak, and A. P. Vogler. 2007. A comprehensive phylogeny of beetles reveals the evolutionary origins of a superradiation. Science 318: 1913-1916. Jungst, S. E., D. R. Farrar, and M. Brandrup. 1998. Iowa’s changing forest resources. Journal of the Iowa Academy of Sciences 105(2): 61-66. 2013 THE GREAT LAKES ENTOMOLOGIST 131

Kasap, H., and R. A. Crowson. 1975. A comparative study of Elateriformia and Das- cilloidea (Coleoptera). Transactions of the Royal Entomological Society of London 126(4): 441-495. King, I. N. 1914. The Coleoptera of Henry County, Iowa. Proceedings of the Iowa Academy of Science 21: 317-340. Kirk, V. M. 1969. A list of the beetles of South Carolina. Part 1 - Northern Coastal Plain. South Carolina Agricultural Experimental Station Technical Bulletin 1033: 1-124. Lawrence, J. F., A. M. Hastings, M. J. Dallwitz, T. A. Paine, and E. J. Zurcher. 2000 onwards. Elateriformia (Coleoptera): Callirhipidae: descriptions, illustrations, identification, and information retrieval for families and subfamilies. Ver- sion: 9 October 2005. Available from http://delta-intkey.com/elateria/www/call.htm (accessed July 2011). Lawrence, J. F., and A. F. Newton, Jr. 1982. Evolution and classification of beetles. Annual Review of Ecology and Systematics 13: 261-290. Lawrence, J. F., and A. F. Newton, Jr. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names), pp 779- 1006. In J. Pakaluk and S. A. Slipinski (eds), Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii, Polska Academia Nauk, Warszawa. (LSAM) Louisiana State Arthropod Museum. 2009. Checklist of the Callirhipidae (Coleoptera) of Louisiana: Zenoa picea: Distribution. Louisiana State Arthropod Mu- seum, Baton Rouge. Available from http://entomology.lsu.edu/coleopteraoflouisiana/ callirhipidae.htm (accessed April 2011). Merrill, E. D. 1937. Palisot de Beauvois as an overlooked American botanist. Proceed- ings of the American Philosophical Society 76: 899-920. Muona, J. 2002. Family 56. Eucnemidae Eschscholtz 1829, pp 152-157. In R. H. Arnett, Jr., M. C. Thomas, P. E. Skelley, and J. H. Frank (eds), American Beetles. Polyphaga: Scarabaeoidea through Curculionoidea, Volume 2. CRC Press, Boca Raton, Florida, USA. Mutel, C. F. 1989. Fragile Giants: A Natural History of the Loess Hills. University of Iowa Press, Iowa City, 304 pp. Mutel, C. F. 2008. The Emerald Horizon: The History of Nature in Iowa. University of Iowa Press, Iowa City, 328 pp. (NCSU) North Carolina State University. 2011. North Carolina State University Insect Collection. Available from http://inventory.ent.ncsu.edu/forms/index.cfm (accessed March 2011). Osten Sacken, B. R. 1862. Descriptions of some larva of North American Coleoptera. Proceedings of the Entomological Society of Philadelphia 1(5): 105-130. Available from http://www.biodiversitylibrary.org/item/22852 (accessed July 2011). Packard, A. S., Jr. 1889. Guide to the study of insects and a treatise on those injurious and beneficial to crops: for the use of colleges, farm-schools, and agriculturists. Henry Holt and Co., New York, 715 pp. Available from http://www.biodiversitylibrary.org/ item/37013 (accessed July 2011). Palisot de Beauvois, A. M. F. J. 1805. Insectes recuellis en Afrique et en Ameriqué dans les royaumes d’Oware et de Benin, a Saint-Domingue et dans les Etats Unis, pendent les anne 1786-1797. First livrasion, pages 6-8, plus Plate VII, Figure I. Paris. Popenoe, E. A. 1877. A list of Kansas Coleoptera. Transactions of the Kansas Academy of Science 5: 21-40. Available from http://www.biodiversitylibrary.org/item/35090 (accessed July 2011). 132 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Say, T. 1835. Descriptions of North American coleopterous insects and observations on some already described. Boston Journal of Natural History 1: 151-203. Available from http://biodiversitylibrary.org/item/100882#page/154/mode/1up (accessed July 2011). Schlicht, D. W., J. C. Downey, and J. C. Nekola. 2007. The Butterflies of Iowa. University of Iowa Press, Iowa City, 233 pp. Schwert, D. P. 1996. Effect of Euro-American settlement on an insect fauna: paleonto- logical analysis of the recent chitin record of beetles (Coleoptera) from Northeastern Iowa. Annals of the Entomological Society of America 89(1): 53-63. Shockley, F. W., and A. R. Cline. 2004. A contribution to the inventory of Coleoptera of Missouri: New records from Benton County. Journal of the Entomological Society of Kansas 77(3): 280-285. Smith, D. D. 1998. Iowa Prairie: original extent and loss, preservation and recovery attempts. Journal of the Iowa Academy of Science 105(3): 94-108. Sommers, S. V. 1874. Catalogue of the Coleoptera from the region of Lake Pontchartrain, La. Bulletin of the Buffalo Society of Natural Sciences 2: 78-99. Available from http:// www.biodiversitylibrary.org/page/5000673 (accessed July 2011). Staines, C. L., Jr. 1983. The Rhipiceridae of Maryland. The Maryland Entomologist 2(2): 38-40. Thomas, D. 2009. A. M. F. J. Palisot de Beauvois 1752-1820. Scarab Workers: World Directory. Available from http://www.museum.unl.edu/research/entomology/workers/ APalisot.htm (accessed July 2011). Turnbull, A. L. 1980. Man and insects: The influence of human settlement on the insect fauna of Canada. The Canadian Entomologist 112(11): 1177-1184. Ulke, H. 1903. A list of the beetles of the District of Columbia. Proceedings of the United States National Museum 25(1275): 1-57. Available from http://www.biodiversitylibrary.org/item/32399 (accessed July 2011). Wickham, H. F. 1911. A list of the Coleoptera of Iowa. Bulletin State University of Iowa Laboratories in Natural History 6(2): 1-40. Weiss, H. 1916. Additions to insects of New Jersey, No. 3. Entomological News and Proceedings of the Entomological Section of the Academy of Natural Sciences of Phila- delphia 27(1): 9-13. Available from http://www.biodiversitylibrary.org/item/20187 (accessed July 2011). Wolf, R. C. 1991. Iowa’s State Parks: Also Forests, Recreation Areas, and Preserves. Iowa State Press, Ames, 224 pp. Young, D. K. 2002. Family 52. Callirhipidae Emden 1924, pp 144-145. In R. H. Ar- nett, Jr., M. C. Thomas, P. E. Skelley, and J. H. Frank (eds). American Beetles. Polyphaga: Scarabaeoidea through Curculionoidea. Volume 2. CRC Press, Boca Raton, Florida, USA. 2013 THE GREAT LAKES ENTOMOLOGIST 133 The Horsehair Worm Gordionus violaceus (Nematomorpha: Gordiida) in Minnesota Philip A. Cochran1, Jacob D. Zanon1, Ben Hanelt2

Abstract A sample of 21 Gordionus violaceus (Baird) (Nematomorpha: Gordioi- dea) was obtained from a puddle in Winona County, southeastern Minnesota. These are the first of this species reported from the upper Midwest. Sex ratio was not significantly different from 50:50 and females and males did not differ significantly in length, but females were of significantly greater diameter than males.

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Three species of horsehair worms (Nematomorpha: Gordioidea) have been reported previously from Minnesota: Gordius robustus Leidy, G. difficilis Montgomery, and Paragordius varius (Leidy) (Cochran et al. 1999, 2004; Martin and Cochran 2005; Cochran 2007). We report herein the collection of a fourth species, Gordionus violaceus (Baird, 1853). According to Schmidt-Rhaesa et al. (2003), G. violaceus has been collected not only in Europe, but also in the northeastern (Massachusetts), central (Missouri and Nebraska), and southwestern (Arizona and California) United States, but it has not been reported from the upper Midwest. Horsehair worms were collected on 8 May 2008 in a puddle (~15 m long × 3-5 m wide × 5 cm deep) on a gravel road, Fairwater Drive, ~ 1 km west of Elba, Winona County, Minnesota. At this point the road skirted the North Branch Whitewater River and floodplain forest on one side and a recently cultivated field on the other side. The horsehair worms were identified as G. violaceus using light microscopy on the basis of male characteristics listed by Schmidt-Rhaesa et al. (2003), including a bi-lobed posterior, absence of a post-cloacal crescent, and presence of pre-cloacal rows of branched bristles that barely reach the tail lobes. In addition, the worms were light brown in body color with a white anterior tip and a dark collar, and both sexes possessed a single type of areole with numerous bristles in the interareolar spaces (Schmidt-Rhaesa et al. 2003). At least some G. violaceus collected previously have been obtained from temporary aquatic habitats (Smith 1991). The sample of G. violaceus included 12 females and nine males, a sex ratio not significantly different from 50:50 P( = 0.664, binomial test). Hanelt et al. (2005) reviewed sex ratios in horsehair worms and noted that sex ratio may vary during the course of the mating season. The presence of sperm plugs on nine females indicated that mating had occurred recently, although the worms when collected were scattered throughout the puddle and not tightly clumped. Preserved females ranged in length from 82 mm to 199 mm and males ranged in length from 152 mm to 207 mm. Both maximum lengths were greater than those reported by Smith (1991) for 10 North American specimens (182 mm for

1Biology Department, Saint Mary’s University of Minnesota, 700 Terrace Heights, Winona, Minnesota 55987. 2 Department of Biology, 167 Castetter Hall, University of New Mexico, Albuquerque, New Mexico 87131. 134 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 females and 132 mm for males). Mean (± SE) lengths of females (157.3 ± 8.4 mm) and males (175.8 ± 6.8 mm) were not significantly different t( = 1.631, df = 19, P = 0.119). However, mean diameter of females (0.54 ± 0.022 mm) was significantly greater than mean diameter of males (0.47 ± 0.015 mm) (t = 2.170, df = 19, P = 0.043). The diameters observed in this study were consistent with those previously reported (Smith 1991: all < 1 mm; Schmidt-Rhaesa et al. 2003: 0.4-0.7 mm). We are not aware of previous comparisons of length or diameter between females and males of this species.

Acknowledgments The horsehair worms were encountered during field surveys for lampreys funded in part by the Minnesota Department of Natural Resources and facilitated by Liz Harper of that organization. We thank Meghan Jensen for assistance in the laboratory. This work was supported in part by the National Science Foundation, DEB–0950066 to BH. Voucher specimens have been placed in the Museum of Southwestern Biology, Parasitology Division, at the University of New Mexico (MSB:PARA: 1002).

Literature Cited Cochran, P. A. 2007. Secondary predation on the horsehair worm Gordius robustus (Nematomorpha; Gordiida). The Great Lakes Entomologist 40: 80-83. Cochran, P. A., A. P. Kinziger, and W. J. Poly. 1999. Predation on horsehair worms (Phylum Nematomorpha). Journal of Freshwater Ecology 14: 211-218. Cochran, P. A., A. K. Newton, and C. Korte. 2004. Great Gordian knots: sex ratio and sexual size dimorphism in aggregations of horsehair worms (Gordius difficilis). Invertebrate Biology 123: 77-81. Hanelt, B., F. Thomas, and A. Schmidt-Rhaesa. 2005. Biology of the Phylum Nema- tomorpha. Advances in Parasitology 59: 243-305. Martin, W. R., and P.A. Cochran. 2005. Horsehair worms (Phylum Nematomorpha) in Iowa and Minnesota. Journal of the Iowa Academy of Science 112: 66-69. Schmidt-Rhaesa, A., B. Hanelt, and W.K. Reeves. 2003. Redescription and compila- tion of Nearctic freshwater Nematomorpha (Gordiida), with the description of two new species. Proceedings of the Academy of Natural Sciences of Philadelphia 153: 77-117. Smith, D. G. 1991. Observations on the morphology and taxonomy of two Parachordodes species (Nematomorpha, Gordioida, Chordodidae) in southern New England (USA). Journal of Zoology, London 225: 469-480. 2013 THE GREAT LAKES ENTOMOLOGIST 135 First Record of Chrysomya rufifacies (Diptera: Calliphoridae) in Wisconsin Jordan D. Marché II1

Abstract The Hairy Maggot Blow Fly, Chrysomya rufifacies (Macquart) (Diptera: Calliphoridae) is recorded for the first time from Wisconsin. Not a record of establishment, the occurrence of this species is that of a seasonal migrant which nonetheless confirms its expected distribution during late summer and early fall in the western Great Lakes region.

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Female and male specimens of the Hairy Maggot Blow Fly, Chrysomya rufifacies (Macquart) (Diptera: Calliphoridae), were collected on successive days, 2 and 3 October 2012, respectively, at the Town of Oregon Park, Dane County, Wisconsin (approximate coordinates: +42° 54' 10", –89° 25' 16.5"), near the author’s home. This finding represents a new state record for the species. Both individuals were collected with a standard aerial net on late blooms of goldenrod (Solidago spp.) and were presumably attracted by the plant’s abundant pollen. However, no additional specimens were seen or collected, despite further searches over the next few weeks, nor were any found upon groups of asters (Aster spp.) that were blooming concurrently, perhaps because of their much-reduced levels of pollen. No nearby sources of carrion, on which the flies must have developed, were found, either. Specimens were identified according to the keys given in the online Canadian Journal of Arthropod Identification(CJAI 2011). C. rufifacies also possesses a prominent whitish anterior thoracic spiracle. Close-up photographs were then submitted to Phil Pellitteri, Insect Diagnostic Laboratory, Department of Entomology, University of Wisconsin-Madison, who tentatively confirmed the identifications and verified that no previous records or specimens were held in the Wisconsin Insect Research Collection (IRC). The two voucher specimens are housed in the author’s private collection (JDMC). The township park where the specimens were collected consists of a moderate-sized grassy area, including a small playground, that is surrounded by much larger untended fields that support abundant native plants such as milkweed (Asclepias spp.) and goldenrod. On the south side, there is an adjoining woodland (with paved and unpaved trails), while on the north and east sides it borders agricultural row crops. Its western side is terminated by housing de- velopments. Other calliphorid and macro-dipteran species found on goldenrod at this site (either before or after the appearance of C. rufifacies) include the Calliphorinae (Cynomya cadaverina Robineau-Desvoidy); Luciliinae (Lucilia sericata (Meigen) and L. silvarum (Meigen)); Polleniinae (Pollenia sp. cf. rudis (Fabricius)); and Chrysomyinae (Cochliomyia macellaria (Fabricius)); along with Muscidae (Musca autumnalis De Geer) and Tachinidae (Archytas apicifer (Walker)). This list is not exhaustive, however. Chrysomya rufifacies is a non-indigenous calliphorid that is native to Australia and southeastern Asia. Accidentally introduced into Central America in the 1970s, the fly was first recorded in the southern U.S. (Texas) in 1982.

15415 Lost Woods Court, Oregon, Wisconsin 53575 (e-mail: [email protected]). 136 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Over the years, it has spread progressively northward, becoming established in the Knoxville, Tennessee area by 1998 (Shahid et al. 2000). By 2007, abundant specimens had been collected as far north as Indiana (Slone and Gruner 2007), although notably fewer individuals had been taken in southern Ontario (Rosati and VanLaerhoven 2007). Belonging to the old-world screwworm flies (subfamily Chrysomyinae), C. rufifacies can be a vector of myiasis in livestock, although it is principally a carrion feeder. It has been found increasingly useful in forensic entomology, where its well-studied lifecycle depends mainly upon the ambient temperature. But if present in sufficient numbers, larval aggregations may produce metabolic heat in excess of ambient temperatures and thus strive to optimize their thermal environment (Slone and Gruner 2007). Adults may live up to 30 days after eclosion (Baumgartner 1993). Previous research has shown that larvae of C. rufifacies are oftentimes dominant competitors to native calliphorid species, especially the Secondary Screwworm Fly, Cochliomyia macellaria (Fabricius) (Baumgartner 1993). Concerns have been raised that the continued spread of C. rufifacies in North America may threaten not only the viability of C. macellaria, but also disrupt the carrion-insect community as a whole (Rosati and VanLaerhoven 2007). Reflecting its tropical origins,C. rufifacies cannot successfully reproduce when temperatures drop into the range of 9–15 °C (Rosati and VanLaerhoven, 2007). As a result, this species cannot overwinter in the northern regions of the U.S. and southern Canada. It must be noted, however, that larvae and adults of C. rufifacies have been obtained from pigs in almost every year since 1998 as part of the collections made for the Forensic Entomology class taught at Michigan State University (R. W. Merritt, personal communication). But this regular occurrence, like that observed in Indiana, appears more likely due to an annual influx of the species into the Great Lakes region from established populations occurring within the southern states. At this time, the species does not appear to be established in Wisconsin and can be nothing more than a seasonal migrant. This conclusion is supported by the fact that specimens of C. rufifacies have not been collected at these latitudes during the spring or early summer months (Rosati and VanLaerhoven 2007). Instead, they have only been taken during late summer and early fall, after successive generations of flies have spread northward. Appearance of these flies might be attributed to the prolonged period of unusually warm and dry weather experienced across the upper Midwest during the spring and summer of 2012. However, the current distributional range of C. rufifacies was expected to include most/all of southern Wisconsin (Fig. 1a, Rosati and VanLaerhoven 2007). Capture of the two Wisconsin specimens thus confirms the expected range of C. rufifacies, and may bolster projections of its future distribution under the assumption of anticipated global warming (Fig. 1b, Rosati and VanLaerhoven 2007). The latter implies both a northward increase of the zone of its permanent establishment in the southern U.S., along with its seasonal migration into the northern U.S. and southern Canada.

Acknowledgments I wish to thank Phil Pellitteri, Insect Diagnostic Laboratory, Department of Entomology, University of Wisconsin-Madison, for tentatively confirming my identifications and verifying that no previous records or specimens ofC. rufifacies were known from the state. Richard W. Merritt, the referee, shared unpublished data concerning the occurrence of C. rufifacies in the East Lansing, Michigan area and furnished additional key references. 2013 THE GREAT LAKES ENTOMOLOGIST 137

Literature Cited Baumgartner, D. L. 1993. Review of Chrysomya rufifacies (Diptera: Calliphoridae). Journal of Medical Entomology 30: 338-352. (CJAI) Canadian Journal of Arthropod Identification (2011). Key to the Subfamilies and Genera of Calliphoridae. Available from http://www.biology.ualberta.ca/bsc/ejournal/mwr_11/mwr_117.htm (accessed 30 Oct. 2011), Key to Eastern Canadian Species in the Subfamily Chrysomyinae. Available from http://www.biology.ualberta.ca/bsc/ ejournal/mwr_11/mwr_1114.htm (accessed 30 Oct. 2011), and Chrysomya rufifacies (Macquart). Available from http://www.biology.ualberta.ca/bsc/ejournal/mwr_11/ mwr_1120.htm (accessed 30 Oct. 2011). Rosati, J. Y., and S. L. VanLaerhoven. 2007. New record of Chrysomya rufifacies (Diptera: Calliphoridae) in Canada: Predicted range expansion and potential effects on native species. The Canadian Entomologist 139: 670-677. Shahid, S. A., R. D. Hall, N. H. Haskell, and R. W. Merritt. 2000. Chrysomya rufifacies (Macquart) (Diptera: Calliphoridae) established in the vicinity of Knoxville, Tennessee, USA. Journal of Forensic Sciences 45: 896-897. Slone, D. H., and S. V. Gruner. 2007. Thermoregulation in larval aggregations of carrion-feeding blow flies (Diptera: Calliphoridae). Journal of Medical Entomology 44: 516-523. 138 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Occurrence of Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae) feeding on Cirsium pitcheri flowers Jordan M. Marshall1 Diabrotica undecimpunctata howardi Barber (adult = spotted cucumber beetle, immature = southern corn rootworm, Coleoptera: Chrysomelidae) is a polyphagous pest found on a broad range for agricultural plants (Krysan 1986). The larvae are root feeders commonly found on plants in the families Cucurbiaceae (gourds and squashes), Poaceae (grasses, including Zea mays L.), and Fabaceae (legumes, including Medicago sativa L.), and can have significant impact on the crop yield depending on the agricultural system (Campbell and Emery 1967, Brust and House 1990). As an adult, D. u. howardi is among the most commonly encountered insects on gourd species (Cucurbita spp. and Lagenaria spp.) (Fronk and Slater 1956). Foliar damage by D. u. howardi can result in significant reduction in crop yield and floral damage includes feeding on anthers and filaments, reducing subsequent pollen production (Brewer et al. 1987, Sasu et al. 2010). Cirsium pitcheri (Torr. ex Eaton) Torr. and A. Gray (Pitcher’s thistle, Asteraceae) is a federally threatened plant species found in open sand dune ecosystems ranging along Lakes Huron, Michigan, and Superior (Voss 1996, Higman and Penskar 1999). C. pitcheri is a monocarpic species (senescing after a single flowering event) with an extended time period between seedling establishment and flowering (5-8 years) (US Fish and Wildlife Service 2002). While habitat protection, which has been implemented in numerous locations within the range of C. pitcheri, is essential for population safety, without adequate reproduction populations will not be maintained or grow. The combination of pollinator efficiency and animal seed predation has significant impact on the final seed crop for a population ofC. pitcheri (Loveless 1984). On 21 and 22 May 2012, I conducted a survey of insect visitors to C. pitcheri individuals in Indiana Dunes National Lakeshore (IDNL) and Indiana Dunes State Park (IDSP). I identified 20 individual C. pitcheri plants (10 in IDNL and 10 in IDSP), observed each plant for 10 minute observation periods on both survey days, and identified floral visitors to family (Table 1). Due to the rarity of C. pitcheri, the selection protocol was not random and those selected for observation included all flowering plants I encountered through a stochastic search of IDNL and IDSP. A subsequent grid survey at both parks (20 m2 circu- lar plots at 50 m spacing with 1 m wide belt transects between plots) resulted in only one additional floweringC. pitcheri individual at IDNL, which was not included in the observations. C. pitcheri plants observed at IDNL were 40.2 m apart (SD 3.6) and those at IDSP were 134.9 m apart (SD 12.3). I found one occurrence of D. u. howardi at IDNL. At IDSP, I observed ten D. u. howardi individuals feeding on five separate C. pitcheri plants (Fig. 1). Two plants had three individuals feeding while another had two individuals feeding. Those C. pitcheri plants with observed D. u. howardi individuals were 100.2 m apart (SD 22.5), with the closest two plants only 1.8 m from each other and with only one occurrence each of D. u. howardi. Those individuals observed were adults that had overwintered and likely were in a time period of active feeding and mating (Krysan 1986). While D. u. howardi is often characterized as an agricultural pest, especially on Cucurbiaceae species, adults do feed on a wide range of other plants,

1Department of Biology, Indiana University-Purdue University Fort Wayne, Fort Wayne, IN 46805. (e-mail: [email protected]). 2013 THE GREAT LAKES ENTOMOLOGIST 139

Table 1. Insect families observed visiting Cirsium pitcheri flowers in Indiana Dunes National Lakeshore (IDNL) and Indiana Dunes State Park (IDSP), including numbers of plants observed visited by a family and individuals in a family observed.

IDNL IDSP Family Plants Visits Plants Visits Coleoptera Chrysomelidae1 1 1 6 7 Curculionidae 0 0 2 4 Diptera Calliphoridae 0 0 1 1 Syrphidae 4 4 10 32 Tephritidae 10 32 8 14 Cecidomyiidae 5 11 4 10 Hymenoptera Apidae 9 32 1 1 Halictidae 0 0 8 15 1Excludes Diabrotica undecimpunctata howardi.

Figure 1. Example of Diabrotica undecimpunctata howardi feeding (circled) in a Cirsium pitcheri flower at Indiana Dunes State Park. 140 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 including numerous species in Asteraceae. Powell et al. (1996) found D. u. howardi commonly in musk thistle flowers (Carduus nutans L., Asteraceae). It has been observed feeding on the leaves of Solidago canadensis L. and S. fistulosa Miller (Asteraceae) (Fontes et al. 1994). Campbell and Meinke (2006) also found D. u. howardi feeding on the flowers of ten different Asteraceae species, including a Cirsium species. However, reductions in pollen production due to adult D. u. howardi floral feeding in non-agricultural species have not been quantified. Invasive species management is important to improving pollination success for C. pitcheri (Baskett et al. 2011). However, other aspects of habitat protection for C. pitcheri may limit D. u. howardi impacts. Snyder and Wise (2000) demonstrated that D. u. howardi exhibited reduced feeding as an antipredator behavior in the presence of a wolf spider, Hogna helluo (Walckenaer) (Araneae: Lycosidae). As a common early successional species, H. helluo may benefit from conservation biological control practices resulting from habitat protection for C. pitcheri in sand dune ecosystems due to the simplicity of the vegetative structure and regular disturbance (Marshall and Rypstra 1999). There is a need to identify if D. u. howardi larvae feed on C. pitcheri roots. However, due to low soil moisture content within dune soils, like those found at IDNL and IDSP, egg survival and larval development may be adequately hindered (Lummus et al. 1983, Brust and House 1990). While survival of immature D. u. howardi poten- tially could be limited in C. pitcheri habitat, adult migration may be continuous from source populations in local agricultural fields (Lawrence and Bach 1989). Feeding impacts by D. u. howardi on C. pitcheri flowering, pollination, and seed development may be important and need to be quantified.

Acknowledgments Insect surveys were conducted under Indiana Department of Natural Re- sources Research and Collecting Permit # NP12-28 and National Park Service Scientific Research and Collecting Permit # INDU-2012-SCI-0007, and funded in part by an IPFW Summer Faculty Grant. I would like to thank the anonymous reviewer for their constructive comments.

Literature Cited Baskett, C. A., S. M. Emery, and J. A. Rudgers. 2011. Pollinator visits to threatened species are restored following invasive plant removal. International Journal of Plant Sciences 172: 411-422. Brewer, M. J., R. N. Story, and V. L. Wright. 1987. Development of summer squash seedlings damaged by striped and spotted cucumber beetles (Coleoptera: Chrysome- lidae). Journal of Economic Entomology 80: 1004-1009. Brust, G. E., and G. J. House. 1990. Effects of soil moisture, no-tillage and predators on southern corn rootworm (Diabrotica undecimpunctata howardi) survival in corn agroecosystems. Agricultural Ecosystems and Environment 31: 199-215. Campbell, L. A., and L. J. Meinke. 2006. Seasonality and adult habitat use by four Diabrotica species at prairie-corn interfaces. Environmental Entomology 35: 922-936. Campbell, W. V., and D. A. Emery. 1967. Some environmental factors affecting feeding, oviposition, and survival of the southern corn rootworm. Journal of Economic Entomology 60: 1675-1678. Fontes, E. M. G., D. H. Habeck, and F. Slansky, Jr. 1994. Phytophagous insects associated with goldenrods (Solidago spp.) in Gainsville, Florida. Florida Entomolo- gist 77: 209-221. Fronk, W. D., and J. A. Slater. 1956. Insect fauna of cucurbit flowers. Journal of the Kansas Entomological Society 29: 141-145. 2013 THE GREAT LAKES ENTOMOLOGIST 141

Higman, P. J., and M. R. Penskar. 1999. Special Plant Abstract for Cirsium pitcheri. Michigan Natural Features Inventory, Lansing, MI, 3 pp. Krysan, J. L. 1986. Biology, distribution and identification of pestDiabrotica , pp. 1-24. In J. L. Krysan and T. A. Miller (eds.). Methods for the Study of Pest Diabrotica. Springer, New York. Lawrence, W. S., and C. E. Bach. 1989. Chrysomelid beetle movements in relation to host-plant size and surrounding non-host vegetation. Ecology 70: 1679-1690. Loveless, M. D. 1984. Population biology and genetic organization in Cirsium pitcheri, an endemic thistle. Ph.D. Dissertation, University of Kansas, Lawrence, KS. Lummus, P. F., J. C. Smith, and N. L. Powell. 1983. Soil moisture and texture effects on survival of immature southern corn rootworms, Diabrotica undecimpunctata howardi Barber (Coleoptera: Chrysomelidae). Environmental Entomology 12: 1529-1531. Marshall, S. D., and A. L. Rypstra. 1999. Spider competition in structurally simple ecosystems. Journal of Arachnology 27: 343-350. Powell, S. D., J. F. Grant, and P. L. Lambdin. 1996. Incidence of above-ground arthropod species on musk thistle in Tennessee. Journal of Agricultural Entomology 13: 17-28. Sasu, M. A., I. Seidl-Adams, K. Wall, J. A. Winsor, and A. G. Stephenson. 2010. Floral transmission of Erwinia tracheiphila by cucumber beetles in a wild Cucurbita pepo. Envionmental Entomology 39: 140-148. Snyder, W. E., and D. H. Wise. 2000. Antipredator behavior of spotted cucumber beetles (Coleoptera: Chrysomelidae) in response to predators that pose varying risks. Environmental Entomology 29: 35-42. US Fish and Wildlife Service. 2002. Pitcher’s Thistle (Cirsium pitcheri) Recovery Plan. Fort Snelling, Minnesota. vii + 92 pp. Voss, E. G. 1996. Michigan Flora. Part III. Dicots (Pyrolaceae-Compositae). The Cran- brook Institute of Science Bulletin 61 and University of Michigan Herbarium, Ann Arbor, MI. 142 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2 Demonstration of Sex Pheromones in Anabolia bimaculata, Hydatophylax argus, and Nemotaulius hostilis (Trichoptera: Limnephilidae) David C. Houghton1

Abstract The presence of extractable semiochemicals for mate attraction is demonstrated experimentally for the first time in Anabolia bimaculata (Walker), Hydatophylax argus (Harris), and Nemotaulius hostilis (Hagen), three species of limnephilid caddisfly. Each species is also the first member of its respective genus to demonstrate pheromone presence. Sex pheromones appear to be widespread within the Trichoptera, although < 1% of species have been tested and there remains substantial research still to be done on pheromone demonstration, chemical composition, and release behavior.

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This note describes the recent discovery of female-produced sex pheromones attracting males within three caddisfly species of the family Limnephilidae—Anabolia bimaculata (Walker), Hydatophylax argus (Har- ris), and Nemotaulius hostilis (Hagen)—based on field trials conducted in northern Lower Michigan in 2011–2012. Pheromones are known to be part of the mating systems of eight insect orders (Pherobase 2012), including 40 caddisfly species within 15 families (Houghton et al. 2009 and references therein). Although much of the recent work on caddisfly pheromones involves physiological and chemical studies (Ivanov and Löfstedt 1999; Bergmann et al. 2001, 2002, 2004; Löfstedt et al. 2008), basic pheromone biology is still in its relative infancy, as their presence has been demonstrated in < 1% of the described caddisfly fauna. Adults of N. hostilis and A. bimaculata were collected in 2011 from ultraviolet lights placed near Fairbanks Creek (N 44.04°, W 85.64°) in late May and early June, respectively. Previous research (Houghton 2002) has suggested no significant difference in pheromonal attractiveness between virgin and post- oviposition females, thus the status of females was not checked. Fifth instar larvae of H. argus were collected in mid May 2012 from the Little Manistee River (N 44.03°, W 85.73°) and reared to adult in a Frigid Units Living Stream™ set to ambient temperature and photoperiod. Adults emerged in late May and were maintained in the laboratory until enough specimens for a trial had emerged (< 72 h). Following the procedure outlined by Wood and Resh (1984), individual specimens of each species were placed into individual 12 ml glass vials with 2–3 ml of HPLC-grade dichloromethane (Fisher Scientific, Pittsburg, PA) for 1–2 hours. Foam Drosophila vial plugs (Carolina Biological, Burlington, NC) were infused with the 2–3 ml extract from each individual specimen (1 female equivalent) and placed into a Trécé Pherocon 1C wing trap (Gempler’s, Belleville, WI). All pheromone trapping occurred at Fairbanks Creek. This stream was the natal habitat for the extracted adults of A. bimaculata and N. hostilis, and

1Department of Biology, Hillsdale College, 33 East College Street, Hillsdale, MI 49242 (e-mail: [email protected]). 2013 THE GREAT LAKES ENTOMOLOGIST 143 was approximately 12 km from the natal habitat for the H. argus specimens. Populations of H. argus were previously known to exist in Fairbanks Creek. Extracts of males, females, and dichloromethane controls were all tested for their ability to attract conspecific males. In all trials, traps were hung in random order from riparian vegetation approximately 1–2 m above the water, and 10 m apart from each other. Each trial began at dusk and lasted 24 hours. Voucher specimens of all tested species have been deposited in the Hillsdale College insect collection. For all three species, the number of conspecific males caught in traps baited with female extracts was significantly higher than the number caught in control traps or those baited with male extracts (Table 1), demonstrating the presence of an extractable female semiochemical for male attraction. Each species is the first within its respective genus shown to use sex pheromones. Seventeen other limnephilid species, however, have previously been documented to use them. Several species in the genus Pycnopsyche, which is the sister genus to Hydatophylax, have been well-studied in their pheromone use (Houghton 2002, Houghton et al. 2009). Likewise, several species in the limnephilid tribe Limnephilini—which includes the genera Anabolia and Nemotaulius—have been demonstrated to use pheromones (Pherobase 2012). It is likely that pheromone use in the Trichoptera is more prevalent than is known, especially when considering the common and widespread use in their sister taxon, the Lepidoptera. Thus, basic caddisfly pheromone demonstration studies remain important.

Acknowledgments I thank my 2012 field ecology class for helping to collectH. argus larvae, and Ashley Logan, Victoria McCaffrey, Kristin Prol, and Angie Pytel for helping to maintain the specimens in the laboratory. The valuable comments of Christer Löfstedt improved an earlier version of the manuscript. Research funding was provided by the Hillsdale College Biology Department. This is paper #4 of the G.H. Gordon BioStation Research Series.

Table 1. Mean (± SD) number of conspecific males caught in traps baited with various extracts from field trials. Each series of three extracts constitutes a separate experiment. Superscript numbers denote statistically distinct groups of means (One-way Analysis of Variance on ranked data with post-hoc Tukey test).

Date Extract n # caught df F p

May 2011 Dichloromethane only 3 0.0 (± 0.0)A N. hostilis males 3 0.0 (± 0.0)A N. hostilis females 4 2.75 (± 0.96)B Summary data: 2 56.5 < 0.001

June 2011 Dichloromethane only 3 0.0 (± 0.0)A A. bimaculata males 4 0.0 (± 0.0)A A. bimaculata females 5 2.4 (± 1.82)B Summary data: 2 9.27 0.007

June 2012 Dichloromethane only 3 0.0 (± 0.0)A H. argus males 4 0.0 (± 0.0)A H. argus females 3 3.33 (± 0.58)B Summary data: 2 122.5 < 0.001 144 THE GREAT LAKES ENTOMOLOGIST Vol. 46, Nos. 1 - 2

Literature cited Bergmann, J., C. Löfstedt, V. D. Ivanov, and W. Francke. 2001. Identification and assignment of the absolute configuration of biologically active methyl-branched ketones from limnephilid caddis flies. European Journal of Organic Chemistry 16: 3175-3179. Bergmann, J., C. Löfstedt, V. D. Ivanov, and W. Francke. 2002. Electrophysi- ologically active volatile compounds from six species of caddisflies. Nova Supplement Entomologia 15: 37–46. Bergmann, J., C. Löfstedt, V. D. Ivanov, and W. Francke. 2004. Identification and synthesis of new bicyclic acetals from caddisflies (Trichoptera). Tetrahedron Letters 45: 3669-3672. Houghton, D. C. 2002. Pheromone use in Pycnopsyche guttifer (Walker) and P. lepida (Hagen) (Trichoptera: Limnephilidae): evidence for pheromonal dialects, pp. 47–54. In W. Mey (ed) Proceedings of the 10th International Symposium on Trichoptera. Nova Supplement Entomologia 15. Houghton, D.C., C. Lux, and M. Spahr. 2009. Demonstration of sex pheromones in Molanna uniophila (Trichoptera: Molannidae), Platycentropus radiatus, Pycnopsyche indiana, and P. subfasciata (Trichoptera: Limnephilidae), with an assessment of in- terspecific attraction between four sympatricPycnopsyche species. The Great Lakes Entomologist 42: 72–79. Ivanov, V. D., and C. Löfstedt. 1999. Pheromones in caddisflies, pp. 149–156. In H. Malicky and P. Chantarmongkol (eds): Proceedings of the 9th International Sym- posium on Trichoptera, Chiang Mai, 1998. Faculty of Science, University of Chiang Mai, Thailand. Löfstedt, C., J. Bergmann, W. Francke, E. Jirle, B. S. Hansson, and V. D. Ivanov. 2008. Identification of a sex pheromone produced by sternal glands in females of the caddisflyMolanna angustata Curtis. Journal of Chemical Ecology 34: 220–228. Pherobase 2012. The pherobase: database of pheromones and semiochemicals. Avail- able from http://www.pherobase.com/ (accessed 22 June 2012). Wood, J. R., and V. H. Resh. 1984. Demonstration of sex pheromones in caddisflies (Trichoptera). Journal of Chemical Ecology 10: 171–175.