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Habitat Associations of in the Ozark Mountains of Author(s): L. E. Dodd, M. J. Lacki, and L. K. Rieske Source: Journal of the Entomological Society, 84(4):271-284. 2011. Published By: Kansas Entomological Society DOI: http://dx.doi.org/10.2317/JKES110324.1 URL: http://www.bioone.org/doi/full/10.2317/JKES110324.1

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 84(4), 2011, pp. 271–284 Habitat Associations of Lepidoptera in the Ozark Mountains of Arkansas

1 1 2 L. E. DODD, M. J. LACKI, AND L. K. RIESKE

ABSTRACT: Lepidoptera were surveyed using blacklight traps during the growing seasons of 2004 and 2005 in two counties that differed in land use patterns in the Ozark Mountains, Arkansas. Marion County is a fragmented landscape; habitats surveyed were upland forest, riparian forest, edge, and field. Crawford County lies in the Ozark National Forest; habitats surveyed were saw, pole, and sapling size classes of timber. $20 mm in wingspan were identified and enumerated. A total of 8326 moths of $324 and 22 families were identified and tabulated. The total number of species and -level composition varied little between landscapes, but the relative occurrence of species varied between landscapes and across habitats. Dominance of common species varied between landscapes with fewer species forming the bulk of the assemblage in the fragmented landscape. Endemism of certain moths in riparian forest suggests this habitat supports many species. In contrast, few species were recorded in field habitats. These data demonstrate the importance of forest habitats for many species in the Ozark Mountains. Our study forms a foundation for understanding species richness patterns of Lepidoptera in the hardwood forests of central . KEY WORDS: Disturbance, diversity, land use, species checklist, species distributions, species richness, species survey

Introduction Lepidoptera are one of the most hyperdiverse assemblages in the forests of North America (Hodges et al., 1983; Covell, 2005). Despite their widespread occurrence, species within this assemblage vary with forest condition (Summerville and Crist, 2008), suggesting that sensitive taxa may serve as indicators of forest quality or health (Summerville et al., 2004). The distribution and knowledge of habitat affiliations of Lepidoptera are broadly known across eastern North America. Data. exist for many areas in central and eastern North America such as the hardwood and conifer forests of (Rings and Metzler, 1988; Rings and Metzler, 1989; Rings et al., 1991; Summerville et al., 1999; Summerville and Crist, 2001; Coleman et al., 2004), the prairies of Kansas (Wright et al., 2003) and the mixed forests of (Landau and Prowell, 1999a, b), but knowledge regarding species presence and composition of the lepidopteran assemblage is lacking for the Ozark Mountains (but see Spencer, 2006). We conducted blacklight trap surveys in the Ozark Mountains of northern Arkansas over two years to document the relative occurrence of species across habitats in two distinct forest landscapes. While the Ozark Mountains are a biodiversity hotspot for flora and fauna (Frazer et al., 1991), this region has a long history of anthropogenic disturbance (Chapman et al., 2006). By addressing the paucity of data that exists for lepidopteran assemblages in this region, we are not only bolstering our knowledge of regional species distributions in North America but

1 Department of Forestry, University of , Lexington, KY 40546 USA. 2 Department of Entomology, University of Kentucky, Lexington, KY 40546 USA. Accepted 11 October 2011; Revised 21 October 2011 E 2011 Kansas Entomological Society 272 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY also providing data pertaining to the influence of general land use patterns on the taxonomic composition of these ecologically-important .

Materials and Methods Study Sites Surveys were conducted at two landscapes in Crawford County and Marion County, Arkansas (ca. 150 km apart). Forests in both counties consist of upland hardwoods, comprised primarily of an overstory of (Quercus sp.) and (Carya sp.). Due to varied human disturbance, the matrices of habitat within the two landscapes differed. Marion County, hereafter referred to as ‘‘fragmented’’ landscape, is located near the border within the Ozark Highlands region (Omerik’s Level III Ecoregion) (Woods et al., 2004). The area is typical of the Elk River Hills; ridges with elevations of 300–400 m are dissected by small streams, yielding a convoluted landscape of steep valleys separated by narrow ridges. The landscape is impacted by small-scale agricultural use and timber harvests; ownership is predominantly private. Habitats surveyed in the fragmented landscape were: upland forest, riparian forest, edge, and field. ‘‘Edge’’ was defined as the interface of forested and non-forested areas. ‘‘Field’’ habitats were agricultural pastureland consisting of non-native grasses used for grazing and hay production. Riparian habitat was associated with mesic site conditions and, at a minimum, an ephemeral presence of water. Crawford County, hereafter referred to as ‘‘forested’’ landscape, is directly north of the Arkansas River along the border, lying both in the Arkansas Valley and Boston Mountain regions (Omerik’s Level III Ecoregion) (Woods et al., 2004). This landscape lies in the midst of the Boston Mountain Ranger District of the Ozark National Forest. The topography is rugged, with deep hollows and steep benched ridges occurring at elevations of 450–550 m. The landscape is heavily forested, and habitats surveyed were a function of the size class of given stands of timber, including sawtimber (Stand Condition Class 10, ‘‘mature,’’ .30.5 cm diameter at breast height [dbh]), poletimber (Stand Condition Class 11, ‘‘immature,’’ ,20.3 cm dbh), and sapling (Stand Condition Class 13, ‘‘adequately stocked’’) size classes, as defined by the USFS (Silvicultural Practices Handbook: FSH 2471.1 R8).

Sampling and Identification Techniques Lepidoptera were surveyed during the growing seasons of 2004–2005 using 10 W blacklight traps (Universal Light Trap, Bioquip Products, Gardena, CA) (Dodd et al., 2008). Survey nights were fair, with temperatures $16uC at sunset, no precipitation, and low wind. Three replicate survey points were established for each habitat in both forested and fragmented landscapes (n 5 9andn 5 12, respectively); see Dodd et al. (2008) for additional explanation. A single replicate per habitat was considered during a single trap night at a landscape. Survey points were spaced far enough apart to ensure no overlap in effective trapping area (i.e., no two traps were visible from one another due to topographic relief and distances .100 m). Trap nights alternated between forested and fragmented landscapes. In 2004, traps were placed on the ground beginning at sunset and operated for five hours. Survey efforts were expanded in 2005; Lepidoptera were captured throughout an entire night by placing a second adjacent trap at each survey point. The resulting two traps VOLUME 84, ISSUE 4 273 per survey point were operated using timer switches (#2835 BioQuip Products, Gardena, CA). Additionally, traps in 2005 were suspended 2.5 m above ground to increase trap efficacy (Burford et al., 1999). A cotton wad soaked in ethyl acetate was placed in each trap to subdue trapped insects. Following a trap night, specimens were sorted and placed in cold storage (4uC) for identification in the laboratory. Specimens were enumerated and identified to species (or in rare cases) using Covell (2005) and Holland (1903), with family-level following Covell (2005). Representative voucher specimens of selected species were retained in a collection at the University of Kentucky’s Forestry Department. We calculated Jaccard’s and Sørensen’s coefficients of similarity for comparisons between landscapes (Southwood, 1978). Additionally, we used EstimateS (v. 8.2) to generate ICE (Lee and Chao, 1994) and Chao 2 (Chao, 1987) species richness estimations. Estimations were based on 1000 randomizations (Summerville and Crist, 2005). We focused our efforts on macrolepidoptera and those microlepidoptera with wings $20 mm (i.e., some Oecophoridae, Yponomeutidae, Cossidae, Tortricidae, Zygaenidae, Megalopygi- dae, , and Pyralidae); thus, our study is not an exhaustive assessment of the assemblage.

Results Moths were surveyed over six nights from July–August in 2004 with each replicate visited once (n 5 21 trap-nights). Each replicate was visited three times from May– August in 2005, with trapping spanning 24 nights (n 5 161 trap-nights as a result of 7 trap malfunctions). Approximately two trap nights occurred per week in both years; inclement weather did not occur on scheduled survey nights. A total of 10,376 moths were captured over our two year study. Of these, 5059 moths were captured in the forested landscape, and 5317 moths were captured in the fragmented landscape. We were able to identify 8326 (80%) beyond the family level, representing 22 families and $342 species (Table 1). Of these, $280 species were identified from the forested landscape and $270 species were identified from the fragmented landscape. Chao 2 estimates of richness were 345 and 321 species for forested and fragmented landscapes, respectively. The ICE estimates of richness were more conservative, with 319 and 320 species for the forested and fragmented landscapes, respectively. The were the most speciose family recorded in both landscapes, followed distantly by the Geometridae, Arctiidae, , and Pyralidae (Fig. 1). These five families formed .80% of the moths identified in both landscapes. A total of 27 species were ubiquitous and captured across all the habitats in both landscapes (Table 1). The majority of these species were in the Noctuidae (7 species), Pyralidae (5), Arctiidae (4), Notodontidae (3), and Saturniidae (3). These five families formed .80% of the species thus identified. Sørensen’s coefficient revealed 76% overlap in species similarity, whereas Jaccard’s coefficient revealed a 61% overlap between landscapes. Some species were endemic to the forested landscape and to specific habitats within this landscape (Table 1). In total, 73 species were unique to the forested landscape; these were primarily the Noctuidae (39 species), Geometridae (11), Pyralidae (5), and Notodontidae (4). These four families formed .80% of the species 274 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Table 1. A checklist of Lepidoptera collected via blacklight traps in forested and fragmented landscapes in the Ozark Mountains, Arkansas, during the growing seasons of 2004–2005. Values presented are the percent contribution to the entire assemblage within habitats. Asterisks denote capture across all habitats. Other symbols denote endemism within a landscape (Forested {, Fragmented {). Taxa in bold are those accounting for .50% of moths identified.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Apatelodidae Apatelodes torrefacta (J. E. Smith)* 0.4 0.5 0.3 0.1 0.4 0.2 0.4 Olceclostera angelica (Grote) ,0.1 ,0.1 ,0.1 0.2 0.1 0.4 Arctiidae Apantesis sp. (Drury, F., Harris)* 0.5 ,0.1 0.2 ,0.1 0.4 5.8 12.2 Cisseps fulvicollis (Hubner) 0.2 2.6 2.7 Cisthene packardii (Grote) 0.2 0.5 0.4 0.6 Clemensia albata (Packard) 2.0 1.7 0.2 1.1 0.5 0.5 Crambidia sp. (Dyar, Packard)* 2.6 1.3 1.0 1.0 0.8 0.2 0.8 Cycinia inopinatus (Hy. Edwards){ ,0.1 Cycinia tenera (Hubner) 0.1 ,0.1 0.5 0.4 Ecpantheria scribonia (Stoll) 0.6 0.2 0.1 Estigmene acrea (Drury){ 0.2 Euchaetes egle (Drury){ 0.4 ,0.1 phasma (Harvey) ,0.1 0.5 ,0.1 ,0.1 ,0.1 0.1 anna (Grote)* 0.8 0.6 1.1 0.1 0.4 0.3 0.4 Grammia arge (Drury){ 0.1 Grammia figurata (Drury){ 0.3 0.4 Grammia oithona (Strecker){ 0.8 Halysidota tessellaris (J. E. Smith)* 8.1 8.2 5.9 1.7 4.8 3.4 6.3 clymene (Brown) ,0.1 ,0.1 0.3 0.1 0.1 0.8 Haploa contigua (Walker) 0.6 0.5 0.2 ,0.1 Haploa reversa (Stretch) ,0.1 0.2 ,0.1 Holomelina aurantiaca (Hubner){ 0.1 0.4 Holomelina opella (Grote) 1.7 0.3 0.4 0.1 Hyphantria cunea (Drury){ 0.3 0.5 1.0 Hypoprepia fucosa (Hubner) 2.6 10.3 0.2 16.5 12.2 2.7 Hypoprepia miniata (Kirby){ 0.2 Pyrrharctia isabella (J. E. Smith) 0.1 0.1 1.3 4.3 Spilosoma congrua (Walker) 2.8 2.2 0.3 1.7 2.1 1.0 Spilosoma virginica (F.) 0.1 0.3 0.3 0.1 0.5 0.4 Cossidae Prionoxystus robiniae (Peck) 0.2 0.3 0.5 ,0.1 Drepanidae Eudeilinea herminiata (Guene´e) 0.1 0.1 Oreta rosea (Walker) ,0.1 ,0.1 0.2 Epiplemidae Calledapteryx dryopterata (Grote) 0.3 0.7 ,0.1 0.1 Callizzia amorata (Packard){ 0.4 0.3 0.3 Geometridae defectaria (Guene´e) 0.4 0.3 0.4 0.2 0.2 0.4 Anacamptodes ephyraria (Walker) ,0.1 0.5 0.1 0.4 0.1 Antepione thisoaria (Guene´e) ,0.1 ,0.1 0.2 multiferata (Walker){ ,0.1 Besma endropiaria (Grote & Robinson) ,0.1 ,0.1 0.1 Besma quercivoraria (Guene´e) ,0.1 ,0.1 0.2 0.1 Calothysanis amaturaria (Walker){ ,0.1 Cyclophora pendulinaria (Guene´e) 0.3 0.6 ,0.1 0.3 0.1 0.1 Dichordia iridaria (Guene´e) 0.2 ,0.1 Dyspteris abortivaria (He¨rrich-Schaffer) 0.1 ,0.1 ,0.1 Ecliptopera atricolorata (Grote & Robinson) 0.1 0.4 0.2 VOLUME 84, ISSUE 4 275

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Ectropis crepuscularia (Dennis & Schiffermu¨ller) ,0.1 0.5 0.3 ,0.1 0.1 Epimecis hortaria (F.) 0.1 0.5 0.1 0.1 Eubaphe mendica (Walker) 0.1 ,0.1 0.1 0.1 0.1 amoenaria (Guene´e) 0.1 0.2 ,0.1 0.1 Euchlaena irraria (Barnes & McDunnough) ,0.1 0.1 Euchlaena obtusaria (Hubner){ ,0.1 Euchlaena pectinaria (Dennis & Schiffermu¨ller) 0.4 0.5 0.2 0.4 0.6 0.3 Euchlaena tigrinaria (Guene´e) ,0.1 ,0.1 Eulithis diversilineata (Hubner) ,0.1 0.3 ,0.1 0.2 0.1 Eumacaria latiferrugata (Walker){ 0.1 Eupithecia miserulata (Grote) 0.6 1.0 1.6 0.7 1.3 1.1 Eusarca confusaria (Hubner) 0.1 0.3 ,0.1 0.3 Eutrapela clemataria (J. E. Smith) 0.1 ,0.1 0.2 0.2 Exelis pyrolaria (Guene´e){ ,0.1 ,0.1 Glena cribrataria (Guene´e) 0.8 0.3 0.3 0.1 0.1 (Hulst) 1.8 ,0.1 0.5 0.5 0.8 Heliomata cycladata (Grote & Robinson){ 0.3 (Walker){ 0.2 (He¨rrich-Schaffer) 0.2 ,0.1 Hydria prunivorata (Ferguson){ 0.2 Hypagyrtis unipunctata (Haworth) 0.4 1.3 1.2 0.9 1.9 3.1 umbrosaria (Hubner) ,0.1 0.1 Idaea demissaria (Hubner) ,0.1 0.1 Idaea furciferata (Packard) 0.5 0.2 0.3 0.5 0.7 0.2 larvaria (Guene´e){ 0.2 ,0.1 ,0.1 Itame coortaria (Hulst) 0.4 3.0 0.4 ,0.1 Lambdina fervidaria (Hubner)* 1.3 0.8 0.2 6.5 5.7 1.5 5.1 Leptostales rubromarginaria (Packard) 0.8 0.3 0.9 0.1 0.9 1.3 Lomographa vestaliata (Guene´e) 0.5 0.3 3.0 ,0.1 0.7 0.2 unitaria (He¨rrich-Schaffer) 0.1 0.3 0.3 1.6 0.6 0.2 Mellilla xanthometata (Walker) 0.8 ,0.1 ,0.1 Metarranthis angularia (Barnes & McDunnough){ 0.2 Metarranthis hypochraria (He¨rrich-Schaffer){ ,0.1 0.3 ,0.1 Nematocampa limbata (Haworth){ 0.2 Nemoria lixaria (Guene´e) ,0.1 0.1 0.2 0.3 ,0.1 0.1 Orthonama centrostrigaria (Wollaston) ,0.1 0.3 0.3 0.3 0.1 Orthonama obstipata (F.) 0.1 ,0.1 0.1 0.4 Patalene olyzonaria (Walker) 0.5 0.5 0.2 0.2 0.1 Pero hubneraria (Guene´e) 0.3 0.5 2.8 ,0.1 0.4 0.3 Phigalia sp. (Cramer, Hulst, Minot) ,0.1 0.2 ,0.1 alcoolaria (Guene´e)* 0.1 ,0.1 ,0.1 0.2 0.2 0.3 1.2 Plagodis fervidaria (He¨rrich- Schaffer){ 0.7 0.1 0.1 276 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Plagodis phlogosaria (Guene´e) ,0.1 0.2 ,0.1 Pleuropucha insulsaria (Guene´e){ 0.4 Probole amicaria (He¨rrich-Schaffer){ 0.2 ,0.1 0.3 Probole nyssaria (Guene´e){ 0.1 0.3 Prochoerodes transversata (Drury) ,0.1 0.3 0.3 Protitame virginalis (Hulst) ,0.1 0.2 ,0.1 0.2 Protoboarmia porcelaria (Guene´e) 0.2 0.2 ,0.1 ,0.1 0.1 Scopula limboundata (Haworth) 0.4 0.5 ,0.1 ,0.1 0.3 0.3 Semiothisa continuata (Walker){ ,0.1 Semiothisa multilineata (Packard) 0.1 ,0.1 0.2 0.2 0.1 Semiothisa ocellinata (Guene´e) 0.1 0.2 0.5 ,0.1 Semiothisa promiscuata (Ferguson){ 0.4 0.3 0.3 Semiothisa quadrinotaria (He¨rrich-Schaffer) 0.2 ,0.1 Semiothisa transitaria (Walker){ 0.1 0.1 0.1 Tetracis crocallata (Guene´e){ ,0.1 Xanthotype urticaria (Sweet) ,0.1 0.1 Lasiocampidae Malacosoma americanum (F.)* 1.1 1.1 5.2 7.3 8.1 6.8 2.4 Malacosoma disstria (Hubner){ 0.7 0.6 0.8 Limacodidae Adoneta spinuloides (He¨rrich-Schaffer){ 0.2 0.1 0.2 Apoda biguttata (Packard){ ,0.1 Apoda y-inversum (Packard) ,0.1 ,0.1 0.4 0.1 ,0.1 0.1 Euclea delphinii (Boisduval) ,0.1 0.1 (He¨rrich-Schaffer) ,0.1 0.3 0.2 Lithacodes fasciola (He¨rrich-Schaffer) 0.3 0.6 0.9 0.3 0.3 Parasa chloris (He¨rrich-Schaffer){ ,0.1 0.1 0.2 badia (Hubner) ,0.1 0.2 0.2 ,0.1 Tortricidia flexuosa (Grote) ,0.1 0.1 ,0.1 ,0.1 Lymantriidae obliquata (Grote & Robinson) 1.3 0.6 0.7 0.3 0.4 0.1 Dasychira tephra (Hubner){ 0.1 Orgyia leucostigma (J. E. Smith) 0.2 ,0.1 ,0.1 Megalopygidae Lagoa crispata (Packard) 0.4 ,0.1 0.2 0.2 0.1 0.8 Norape ovina (Sepp) 0.8 0.7 1.0 ,0.1 0.2 0.3 Mimallonidae Lacosoma chiridota (Grote) 0.1 ,0.1 ,0.1 0.1 0.4 Noctuidae Abagrotis alternata (Grote){ 0.3 0.1 Acronicta americana (Harris) 0.4 0.2 0.2 Acronicta afflicta (Grote) 0.3 0.3 0.3 0.3 0.3 0.1 Acronicta funeralis (Grote & Robinson){ ,0.1 0.3 Acronicta haesitata (Grote)* 0.3 0.4 1.2 1.5 0.4 0.9 0.4 Acronicta impleta (Walker){ 0.1 ,0.1 0.5 Acronicta inclara (J. B. Smith){ ,0.1 Acronicta interrupta (Guene´e){ 0.1 Acronicta lithospila (Grote){ ,0.1 0.2 Acronicta lobeliae (Guene´e){ 0.1 Acronicta modica (Grote)* 0.2 ,0.1 0.5 0.3 0.1 0.2 0.4 Acronicta noctivaga (Grote){ ,0.1 Acronicta ovata (Grote){ 0.5 0.3 0.3 Acronicta retardata (Walker) ,0.1 0.2 0.3 0.5 0.1 0.1 VOLUME 84, ISSUE 4 277

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Acronicta tritona (Hubner){ ,0.1 Agriopodes fallax (He¨rrich-Schaffer) ,0.1 ,0.1 ,0.1 ,0.1 0.1 Agriopodes teratophora (He¨rrich-Schaffer) 0.1 ,0.1 Agrotis ipsilon (Hufnagel) ,0.1 0.1 ,0.1 ,0.1 0.4 Allagrapha aeria (Hubner) ,0.1 0.4 Allotria elonympha (Hubner) 0.7 0.8 0.6 0.3 0.4 0.5 Amphipyra pyramidoides (Guene´e){ ,0.1 0.1 Anicla infecta (Ochsenheimer){ ,0.1 Anorthodes tarda (Guene´e){ 0.3 anilis (Drury){ 0.1 Autographa biloba (Stephens){ 0.4 australis (Grote){ 0.1 ,0.1 Baileya levitans (J. B. Smith) 2.0 2.8 2.9 ,0.1 0.1 labecula (Grote) 0.4 0.2 0.4 0.3 0.6 0.5 Balsa tristrigella (Walker){ ,0.1 Bleptina caradrinalis (Guene´e) 0.3 ,0.1 0.2 0.1 0.5 0.3 Bomolocha sp. (Grote, Guene´e, Walker){ 0.1 0.1 Bomolocha abalienalis (Walker){ ,0.1 Bomolocha bijugalis (Walker){ ,0.1 Bomolocha sordidula (Grote){ ,0.1 chloropha (Hubner) 0.1 ,0.1 0.3 0.8 Caenurgia erechtea (Cramer) ,0.1 0.9 3.9 Callopistria cordata (Ljungh){ ,0.1 0.1 0.2 Callopistria mollissima (Guene´e){ 0.5 agrippina (Strecker){ ,0.1 Catocala amica (Hubner) 0.1 0.7 Catocala andromedae (Guene´e) ,0.1 ,0.1 0.1 Catocala dejecta (Strecker){ ,0.1 0.4 Catocala epione (Drury){ ,0.1 Catocala flebilis (Grote){ ,0.1 Catocala gracilis (W. Edwards) 0.1 0.1 0.4 Catocala ilia (Cramer) 0.1 0.1 0.3 0.5 0.3 Catocala judith (Strecker){ ,0.1 0.3 Catocala junctura (Walker){ ,0.1 Catocala lacrymosa (Guene´e){ ,0.1 Catocala nebulosa (W. Edwards){ ,0.1 0.1 Catocala neogama (J. E. Smith){ ,0.1 Catocala obscura (Strecker) 0.2 ,0.1 Catocala retecta (Grote){ ,0.1 0.4 Catocala ultronia (Hubner) 0.3 0.1 Catocala vidua (J. E. Smith){ ,0.1 Cerma cerintha (Treitschke) ,0.1 ,0.1 ,0.1 Charadra deridens (Guene´e) ,0.1 Chytonix palliatricula (Guene´e) 0.4 ,0.1 0.4 0.5 0.2 Cosmia calami (Harvey) ,0.1 ,0.1 0.8 0.1 0.3 0.4 festivoides (Guene´e){ 0.1 Elaphria grata (Hubner) 0.1 0.2 ,0.1 0.1 1.2 Elaphria versicolor (Grote) 1.4 0.3 0.1 Eosphoropteryx thyatyroides (Guene´e){ ,0.1 ,0.1 278 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Euagrotis lubricans (Guene´e){ 0.4 Eudryas grata (F.) 0.1 0.5 0.3 ,0.1 Euparthenos nubilis (Hubner){ 0.1 0.2 Euplexia benesimilis (McDunnough){ 0.4 0.6 0.3 Faronta diffusa (Walker) ,0.1 ,0.1 0.3 Galgula partita (Guene´e) 0.4 0.1 0.2 0.1 2.5 3.5 Harrisimemna trisignata (Walker) ,0.1 ,0.1 ,0.1 Heliothis turbatus (Walker){ 0.2 Heliothis zea (Boddie){ ,0.1 Hemeroplanis scopulepes (Haworth){ 0.1 Homohadena badistriga (Grote) ,0.1 ,0.1 Homophoberia apicosa (Haworth){ ,0.1 Hyperstrotia pervertens (Barnes, McDunnough) 0.3 ,0.1 1.0 2.9 2.2 1.4 hormos (Hubner) ,0.1 0.2 0.2 Hypsoropha monilis (F.) ,0.1 0.1 0.1 Idia aemula (Hubner){ ,0.1 Idia americalis (Guene´e)* 1.1 1.3 0.6 1.5 0.7 2.5 0.4 Isogona tenuis (Grote) 0.1 0.3 0.1 Lacinipolia lorea (Guene´e) ,0.1 Lacinipolia renigera (Stephens) 0.6 0.4 0.2 0.4 1.7 1.2 Lesmone detrahens (Walker) 0.1 Leucania sp. (Forbes, Guene´e, Walker){ 0.1 0.1 0.4 Leucania inermis (Forbes){ 0.3 Leucania scirpicola (Guene´e){ 0.1 0.1 2.2 2.4 Lithacodia carneola (Guene´e) 0.8 1.0 1.5 ,0.1 0.3 0.3 Lithacodia muscosula (Guene´e) 1.0 0.5 0.5 ,0.1 2.8 0.1 Macrochilo absorptalis (Walker) ,0.1 0.1 0.4 Meganola minuscula (Zeller){ ,0.1 0.2 Mocis texana (Mo¨rrison) ,0.1 ,0.1 ,0.1 0.1 0.1 Nedra ramosula (Guene´e){ ,0.1 Ogdoconta cinereola (Guene´e)* 0.4 0.3 0.6 0.2 0.2 0.1 0.4 Orthodes crenulata (Butler)* 1.2 ,0.1 0.3 0.2 ,0.1 0.3 0.4 Orthodes cynica (Guene´e) 3.5 ,0.1 0.3 0.1 Ozarba aeria (Grote){ 0.1 abrostoloides (Guene´e) ,0.1 ,0.1 0.2 ,0.1 Paectes oculatrix (Guene´e) 0.1 0.2 ,0.1 Paectes pygmaea (Hubner) 0.1 0.1 ,0.1 ,0.1 0.1 Palthis sp. (Guene´e, Hubner) 0.4 0.3 1.0 ,0.1 0.2 Pangrapta decoralis (Hubner) 0.4 ,0.1 0.7 Panopoda carneicosta (Guene´e) 0.1 ,0.1 0.6 ,0.1 1.2 Panopoda rufimargo (Hubner) 0.2 ,0.1 0.2 0.5 ,0.1 1.2 Panthea furcilla (Packard){ 0.2 0.1 Parallelia bistriaris (Hubner){ ,0.1 Phalaenophana pyramusalis (Walker)* 0.6 0.3 0.9 0.2 0.3 0.8 6.3 Phalaenostola metonalis (Walker){ ,0.1 Phosphila miselioides (Guene´e){ ,0.1 Plathypena scabra (F.) 0.1 ,0.1 0.3 ,0.1 0.4 0.2 Platysenta sutor (Guene´e){ 0.1 ,0.1 Platysenta vecors (Guene´e) 0.1 0.3 0.3 ,0.1 0.2 0.1 VOLUME 84, ISSUE 4 279

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Polia sp. (Grote, Guene´e, Walker) ,0.1 0.2 ,0.1 Polygrammate hebraeicum (Hubner) 3.0 3.1 3.2 2.7 6.2 2.1 Protolampra brunneicollis (Grote) ,0.1 ,0.1 Proxenus miranda (Grote){ 0.8 Pseudaletia unipuncta (Haworth){ 0.1 Pseudeva purpurigera (Walker){ ,0.1 Pseudorthodes vecors (Guene´e){ ,0.1 herbarum (Guene´e){ ,0.1 Rachiplusia ou (Guene´e){ ,0.1 Renia discoloralis (Guene´e) 0.3 0.4 0.2 0.2 0.1 1.6 Renia fraternalis (J. B. Smith) ,0.1 0.2 0.1 0.1 Renia sobrialis (Walker){ 1.1 0.6 0.5 0.4 Renia nemoralis (Barnes & McDunnough) 0.3 ,0.1 Rivula propinqualis (Guene´e){ ,0.1 Schinia arcigera (Guene´e){ 0.2 Schinia lynx (Guene´e){ 0.1 0.9 0.4 Schinia trifascia (Hubner){ 0.2 Scolecocampa liburna (Geyer) ,0.1 0.2 0.1 macula (Druce) ,0.1 0.3 0.2 0.9 0.1 0.2 Spodoptera ornithogalli (Guene´e) 0.4 0.3 ,0.1 ,0.1 Spragueia sp. (Guene´e, He¨rrich-Schaffer){ 0.1 0.1 Stiriodes obtusa (He¨rrich-Schaffer) ,0.1 ,0.1 0.1 0.1 Tarachidia candefacta (Hubner){ ,0.1 0.8 mynesalis (Walker) 3.3 4.2 0.4 11.7 4.5 6.5 Thioptera nigrofimbria (Guene´e)* 2.3 0.8 0.9 0.1 1.9 0.7 1.2 Tricholita signata (Walker){ ,0.1 Trichoplusia ni (Hubner){ ,0.1 Xestia smithii (Snellen){ ,0.1 Zale lunata (Drury) ,0.1 ,0.1 0.1 Zale lunifera (Hubner){ ,0.1 Zanclognatha cruralis (Guene´e) 0.5 0.2 0.2 0.6 0.9 Zanclognatha lituralis (Hubner){ ,0.1 Zanclognatha obscuripennis (Grote) 0.3 0.3 0.3 0.3 Notodontidae Clostera inclusa (Hubner) ,0.1 ,0.1 Datana angusii (Grote & Robinson) 0.3 0.2 0.5 Datana contracta (Walker)* 0.9 0.4 0.3 ,0.1 0.1 0.3 0.4 Datana ministra (Drury) ,0.1 0.1 0.8 Datana perspicua (Grote & Robinson) 0.6 0.8 0.3 0.3 0.1 0.4 biundata (Walker){ 0.1 Heterocampa guttivitta (Walker) 1.6 1.6 0.9 0.4 0.3 Heterocampa obliqua (Packard)* 1.3 1.1 1.0 0.5 0.3 0.6 0.8 Heterocampa subrotata (Harvey) 0.1 0.3 ,0.1 Heterocampa umbrata (Walker){ 0.2 0.2 0.4 georgica (He¨rrich-Schaffer) 0.1 0.3 1.1 ,0.1 0.1 Lochmaeus bilineata (Packard) 0.1 0.3 0.5 0.1 ,0.1 0.5 Lochmaeus manteo (Doubleday){ 0.8 0.3 0.2 Macrurocampa marthesia (Cramer) 0.3 0.7 0.5 ,0.1 0.1 Nadata gibbosa (J. E. Smith)* 0.5 0.7 0.4 1.1 0.7 1.3 0.8 280 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Nirece bidentata (Walker) 0.1 ,0.1 ,0.1 ,0.1 ,0.1 Oligocentria lignicolor (Walker) 0.3 0.3 0.1 Oligocentria semirufescens (Walker){ ,0.1 Peridea angulosa (J. E. Smith){ ,0.1 Peridea basitriens (Walker) ,0.1 0.3 0.4 Schizura ipomeae (Doubleday) 0.3 0.2 0.3 Schizura leptinoides (Grote) ,0.1 0.1 0.4 Symmerista albifrons (J. E. Smith) 0.4 0.1 0.3 0.1 Oecophoridae leucillana (Zeller) 0.3 0.1 0.2 0.1 Antaeotricha schlaegeri (Zeller) ,0.1 0.4 0.1 ,0.1 0.5 Psilocorsis sp. (Clemens) 0.8 0.5 0.4 0.9 0.5 0.3 Pterophoridae Platyptilia carduidactyla (Riley){ ,0.1 Pyralidae ranalis (Guene´e)* 0.3 0.4 1.2 1.9 2.6 1.0 1.6 Clydonopteron tecomae (Riley){ 0.2 capitalis (Grote) 0.2 ,0.1 ovulalis (Guene´e){ 0.3 ,0.1 Crambus agitatellus (Clemens)* 1.8 3.2 1.2 0.3 2.1 3.4 1.2 Crambus laqueatellus (Clemens)* ,0.1 ,0.1 ,0.1 ,0.1 0.2 0.8 0.4 Desmia funeralis (Hubner)* 2.0 1.7 2.3 0.4 0.6 0.9 0.4 elealis (Walker){ ,0.1 ,0.1 Epipagis huronalis (Guene´e){ 0.1 Helvibotys helvialis (Walker) 0.2 0.3 0.2 ,0.1 0.1 Herculia olinalis (Guene´e)* 0.1 0.1 ,0.1 3.4 0.3 1.6 0.4 Hymenia perspectalis (Hubner){ 0.1 Munroessa gyralis (Hulst){ ,0.1 Nomophila nearctica (Munroe){ ,0.1 Palpita magniferalis (Walker) 7.8 6.0 6.6 0.4 2.7 0.7 Pantographa limata (Grote & Robinson) 1.1 0.6 0.3 0.4 flavidalis (Guene´e) ,0.1 0.1 0.5 0.6 1.6 Pyrausta acrionalis (Walker) 0.3 0.3 Tetralopha asperatella (Clemens) 1.3 0.8 1.9 1.3 0.8 0.6 Udea rubigalis (Guene´e) 0.6 0.3 0.3 0.3 1.1 0.4 Urola nivalis (Drury) 0.2 ,0.1 ,0.1 0.3 1.8 2.0 Saturnidae Actias luna (L.)* 1.0 0.8 0.6 ,0.1 0.5 0.5 1.2 Anisota stigma (F.) 0.1 ,0.1 0.1 0.1 Antheraea polyphemus (Cramer) 0.2 0.3 0.2 0.3 Automeris io (F.)* 0.3 0.3 ,0.1 0.1 0.1 0.1 0.8 Citheronia regalis (F.) 0.2 0.4 Dryocampa rubicunda (F.) 1.4 0.9 0.5 0.4 Eacles imperialis (Drury)* 0.2 1.0 0.3 0.3 0.8 0.3 2.0 Sphingicampa bicolor (Harris){ ,0.1 Sphingidae Ceratomia hageni (Grote){ ,0.1 Ceratomia undulosa (Walker){ 0.8 Darapsa myron (Cramer) ,0.1 ,0.1 ,0.1 0.1 0.4 Darapsa pholus (Cramer){ 0.8 Laothe juglandis (J. E. Smith){ 0.2 ,0.1 Manduca sp. (F., Haworth, L.) ,0.1 ,0.1 Paonias excaecatus (J. E. Smith) 0.3 0.2 ,0.1 0.4 Paonias myops (J. E. Smith){ ,0.1 ,0.1 VOLUME 84, ISSUE 4 281

Table 1. Continued.

Forested landscape Fragmented landscape

Family Species Saw Pole Sapling Upland Riparian Edge Field

Thyatiridae Habrosyne scripta (Gosse){ 0.1 Pseudothyatira cymatophoroides (Guene´e){ ,0.1 Tortricidae Archips argyrospila (Walker) 0.6 ,0.1 1.2 1.1 0.7 2.0 Argyrotaenia alisellana (Robinson) 0.5 0.2 ,0.1 0.1 ,0.1 2.7 Argyrotaenia quercifoliana (Fitch){ 0.5 0.2 0.2 0.4 Choristoneura parrallela (Robinson) 0.8 0.7 0.3 0.1 0.4 0.7 Choristoneura pinus (Freeman){ 0.3 Choristoneura rosaceana (Harris)* 0.3 0.6 2.5 0.9 1.4 0.9 1.6 Sparganothis reticulatana (Clemens) 0.2 ,0.1 0.3 Yponomeutidae Atteva punctella (Cramer) 0.8 0.7 2.7 0.2 0.3 2.0 Yponomeuta sp. (Clemens, Dyar) 0.1 ,0.1 ,0.1 0.5 0.1 Zygaenidae Harrisina americana (Guene´e) 0.2 ,0.1 ,0.1 ,0.1 Pyromorpha dimidiata (He¨rrich-Schaffer){ 0.1 thus identified. Habitat specificity was common within the forested landscape; 82 species were found in only a single forested habitat. Of these, 35 species were in sawtimber habitat, followed by sapling and poletimber habitats (29 and 18 species, respectively). Various species were also endemic to the fragmented landscape and to specific habitats within this landscape (Table 1); 61 species were unique to the fragmented landscape. Of these, families with the most species were the Noctuidae (34 species), Geometridae (9) and Arctiidae (6), comprising .80% of the species thus identified. As in the forested landscape, habitat specificity was commonplace in the fragmented landscape; 87 species were found only in a single habitat in the fragmented landscape. Of these, 31 species were in riparian forest, followed by edge, upland forest, and field habitats (22, 21, and 6 species, respectively). The most common species found in this study were present in both landscapes. A total of 20 species/genera formed .50% of the moths identified (Table 1). In the forested landscape 21 species/genera formed .50% of the moths captured. Of these, Halysidota tessellaris J. E. Smith (7.5%)andPalpita magniferalis Walker (6.8%) dominated. The remaining 19 species comprised ,5% of the total: Hypoprepia fucosa Hubner, Polygrammate hebraeicum Hubner, Tetanolita mynesalis Walker, Baileya levitans J. B. Smith, Malacosoma americanum F., Crambus agitatellus Clemens, Desmia funeralis Hubner, Spilosoma congrua Walker, Crambidia sp., Heterocampa guttivitta Walker, Clemensia albata Packard, Itame coortaria Hulst, Thioptera nigrofimbria Guene´e, Tetralopha asperatella Clemens, Atteva punctella Cramer, Lomographa vestaliata Guene´e, Heterocampa obliqua Packard, Pero hubneraria Guene´e, Lithacodia carneola Guene´e. In contrast, only 13 species/genera formed .50% of the assemblage captured in the fragmented landscape. Four species were numerically dominant, including H. fucosa (11.0%), M. americanum (7.2%), T. mynesalis (7.2%), and Lambdina fervidaria Hubner (5.1%). The remaining nine species comprised ,5% of the total: P. hebraeicum, H. tessellaris, Apantesis sp., Hyperstrotia pervertens Barnes & McDunnough, Blepharomastix ranalis Guene´e, 282 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Fig. 1. Percent composition of lepidopteran families collected via blacklight traps in forested and fragmented landscapes in the Ozark Mountains, Arkansas, during the growing seasons of 2004–2005.

Hypagyrtis unipunctata Haworth, C. agitatellus, Herculia olinalis Guene´e, S. congrua. Based on these more common species/genera, Sørensen’s coefficient revealed 41% overlap in similarity between landscapes, whereas Jaccard’s coefficient revealed a 26% overlap.

Discussion Our study demonstrates little difference in lepidopteran family-level composition between fragmented and forested landscapes, but we document varied occurrence of species between landscapes. Our data suggest that species richness is comparable between fragmented and forested landscapes; similar trends for relatively high lepidopteran richness in fragmented land parcels has been documented further east in North America (i.e., Ohio) (Summerville and Crist, 2001). Even so, our data demonstrate that the dominance of common species within families differed between the two landscapes. Summerville, et al. (2008) noted relative differences in the VOLUME 84, ISSUE 4 283 dominance of common species between states and even regions of eastern North America. Our study demonstrates differences in species dominance at a finer scale, likely due to different land use patterns and associated flora in the two landscapes. It is intriguing that we observed fewer species forming the bulk of the assemblage in the fragmented landscape, suggesting that fragmentation and/or agricultural land use holds consequence for species evenness. At a finer resolution, we documented varied occurrence of species across habitats. While endemism across habitats in the forested landscape was similar, more dramatic variation was observed in the fragmented landscape. The number of species found only in riparian areas suggests this habitat is important for lepidopteran species richness. Patterns in vegetation richness, particularly in the understory, likely drive our observations (Ober and Hayes, 2009). In contrast, the relatively few number of species found only in the field habitat suggests this habitat supports fewer moth species than forest habitats; not surprising given the stark homogeneity of vegetation in the field habitat versus the diversity of vegetation found in forested habitats. Our study is not exhaustive and possesses biases inherent in similar studies (Summerville et al., 1999). We only sampled through a portion of the growing season (20 May–11 August), so we are likely missing many early or late-occurring species. Further, we only present data from a single sampling approach, thus our study is biased towards phototaxic taxa. Finally, our study focused on moths with wingspans $20 mm, and emphasized larger taxa. Despite these limitations, we present a working checklist for Crawford and Marion counties in Arkansas with species richness levels comparable to that reported in other checklists for North America (Landau and Prowell, 1999a, b; Summerville et al., 1999; Wright et al., 2003; Coleman et al., 2004). Our species richness estimations suggest that we recorded the vast majority of possible species targeted by our methods ($86% in Crawford County and $84% in Marion County). Biological inventories, particularly species checklists, are an undervalued tool despite their role as a ‘‘first step’’ in implementing sound conservation strategies (Summerville et al., 1999). Given the overlap in species between landscapes, our study presents a foundation for further lepidopteran inventories in the Ozark Mountains and, more broadly, the hardwood forests of central North America.

Acknowledgements This research was made possible through funding provided by the Arkansas Game and Fish Commission, the U.S. Fish and Wildlife Service, and the University of Kentucky’s Department of Forestry. Our study was conducted under agreements with the Arkansas Game and Fish Commission, the Arkansas Natural Heritage Commission, The Nature Conservancy, the U.S. Forest Service, and the U.S. Fish and Wildlife Service. We thank the U.S. Forest Service, The Nature Conservancy, the Arkansas Natural Heritage Commission, and private landowners for access to land holdings. We thank T. Barnes, B. Sasse, and W. Puckette for advice with study design, as well as R. Coy for technical assistance. This is paper 11-09-035 of the Kentucky Agricultural Experiment Station and is published with the approval of the director.

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