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MICROLEPIDOPTERA OF ILLINOIS HILL

BY

TERRY L HARRISON

DISSERTATION

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in in the Graduate College of the University of Illinois at Urbana-Champaign, 2011

Urbana, Illinois

Doctoral Committee:

Professor May R. Berenbaum, Chair and Director of Research Professor Stewart H. Berlocher Professor James B. Whitfield Brenda Molano-Flores, Illinois Natural History Survey

ABSTRACT

Among , which are the most diverse eukaryotic group on earth, is one of four enormously diverse orders, with approximately 10,000 described in North

America. Within the order, Nearctic “,” which represent an overwhelmingly large percentage of diversity within the order, remain poorly known despite their ecological importance in many communities. In this thesis, I undertook several studies of microlepidoptera diversity in a natural community (hill ) and a managed community type (biofuel feedstock). In two Illinois hill prairies differing in size, latitude, and plant composition, alpha diversity of and was similar, but the prairies were found to support different sets of species of these groups. It is concluded that the similarity in alpha diversity occurs because the larger prairie supports primarily a complement of moth species that feed as larvae on prairie (especially species of and ), whereas the collected in the small prairie represent relatively few prairie-associated species, plus a large component of species that feed as larvae on deciduous trees that surround the prairie. This agrees with the finding of high beta diversity of moths between the sites, which reflects a high level of larval hostplant specificity in most species of Pyraloidea and Tortricidae.

Based on published information plus observations made on microlepidoptera collected during the course of this study, 31 families of microlepidoptera were reviewed with an aim toward evaluating the likelihood of their including species that are dependent on tallgrass prairie.

Of these families, 12 were evaluated as possible, and two as likely or certain, to include prairie- dependent species.

ii In a comparison of moth diversity in light-trap samples from corn, miscanthus,

switchgrass, and native prairie, alpha diversity was highest in prairie and was higher in

switchgrass than in the other two biofuel crops. Moth species complements generally were

similar among the biofuel crops, and prairie shared higher species complementarity with

switchgrass than with corn or miscanthus. These findings suggest that large-scale conversion of

land to biofuel crops may, to a substantial degree, detrimentally affect ,

with a resulting loss of valuable arthropod-derived ecosystem services both within the cropping

systems and in the surrounding landscape.

During the course of this study, rearing efforts yielded two species of moths of the

, both of which are monophagous leaf feeders on leadplant, Amorpha

canescens (Fabaceae). Because these moths are restricted to tallgrass prairie, they are likely to

be of interest to conservation biologists. In the interest of naming the moths to facilitate communication regarding them, and to augment our taxonomic knowledge of their respective genera, the moths are described, and diagnoses are provided to differentiate them from similar, related species.

iii ACKNOWLEDGMENTS

I thank the staff of the Illinois Department of Natural Resources and the staffs of the EBI

Energy Farm, Orr Agricultural Research and Demonstration Center, Northern Illinois Agronomy

Research Center, Brownstown Agronomy Research Center, Brownstown Agronomy Research

Center, and Dixon Springs Agricultural Center, for granting me permission to collect moths on lands under their stewardship. A. Abbas, C. Markell, L. Eggers, K. Merriman, A. Yanahan, D.

Pearlstein, R. Orpet, A. Lawrance, and A. Lovdahl are thanked for assistance in collecting and/or processing specimens. A. Wallner is thanked for assistance with data analysis. A. Zangerl is gratefully acknowledged for valuable assistance in many different areas of this project. J. Taft is thanked for providing information on in Illinois. This study was supported in part by grants from Prairie Biotic Research and the Xerxes Society, to whom I am grateful.

Finally, I thank my advisory committee, and especially my major adviser, Dr. May R.

Berenbaum, for the guidance that they provided to me in this study.

iv TABLE OF CONTENTS

Chapter 1. Pyraloidea and Tortricidae (Lepidoptera) in Two Illinois Hill Prairies …………….. 1

Chapter 2. Evaluation of Prairie-Dependent Microlepidoptera of the Basal Families

Micropterigidae through Gelechiidae ……………………………………………………..…… 26

Chapter 3. Moth Diversity in Three Biofuel Crops and Native Prairie ……………………….. 40

Chapter 4. A New, Prairie-Restricted Species of Filatima Busck (Lepidoptera: Gelechiidae) from Illinois ……………………………………………………………….………………….... 65

Chapter 5. A New, Prairie-Restricted Species of Anacampsis Curtis (Lepidoptera: Gelechiidae) from Illinois …………………………………………………………………….……………… 78

Author’s Biography ……………………...………………………………………………..…… 91

v Chapter 1. Pyraloidea and Tortricidae (Lepidoptera) in Two Illinois Hill Prairies

Abstract. I compared diversity, abundance, and species composition of Pyraloidea

(= + sensu Munroe and Solis 1998) and Tortricidae in two Illinois loess hill prairies having similar management history, but differing in size, latitude, and some aspects of plant species composition. A total of 2798 moths representing 214 species of Pyraloidea and

Tortricidae were examined. Alpha-diversity values were similar for the two sites, whether for combined samples or for Pyraloidea or Tortricidae considered separately. Beta diversity, however, was high, indicating dissimilar species compositions in the two prairies, especially for

Tortricidae. Moth species compositions generally reflected high diversity and abundance of

Asteraceae and Fabaceae at Revis, and proximity to forest at Pere Marquette, except that the predominant tortricid species at the latter site was the leadplant-feeding taleana.

No significant differences were seen in moth abundance per species among collection dates, when compared either by site or by taxon. This study demonstrates that even small tracts with comparatively low plant diversityshould not be disregarded in conservation plans, in that they can serve as refugia for some species of prairie-dependent microlepidoptera.

Key Words. Hill prairies, native biodiversity, conservation biology, restricted habitat, moths, microlepidoptera

Introduction

Conservation of biodiversity, in addition to being of aesthetic and scientific interest, is becoming increasingly recognized as an economic concern of global importance. As of June,

1 1997, more than 170 nations had explicitly acknowledged this economic significance by

ratifying the Convention on Biodiversity (Raven et al. 1998). Such acknowledgment seems

appropriate considering that direct benefits of biodiversity can be seen in agriculture, fisheries,

forest products, pharmaceuticals, medical and research tools, nature travel, horticultural trade,

and “ecosystem services”: pollination, seed dispersal, detoxification and water purification,

removal and storage of atmospheric carbon dioxide, grazing, fisheries protection, and flood

control. Information on biodiversity has been developed into important applications such as the breeding of disease resistance from wild plants into related cultivars, and the prolific exploitation of molecular biology techniques, knowledge of which began with biodiversity studies of bacteria

(Raven et al. 1998).

Raven et al. (1998) asserted that the sometimes confrontational relationship between

economy and the environment is in need of realignment through development of a program of

sustainable management of “natural capital.” One of their recommendations for achieving this

goal is the establishment of “an objective, accessible knowledge base that includes what we

know about species, their characteristics and interactions, their habitats and ecosystems, how

human activities impact them, and what kinds of actions comprise best practices for managing

them.”

Current knowledge and understanding of North American biodiversity is too superficial

to allow establishing such a knowledge base. In fact, for many biotic groups, even the most

basic step, mere documenting of species, has never been taken. Raven et al. (1998) stated that

fewer than 30% of species in the USA have been documented and described; they recommended

that $130 million per year in federal funding be allocated solely to discovery of species.

Similarly, Donoghue and Alverson (2000) noted that in most major lineages, the number of new

2 species being described is increasing, and that this trend has resulted not from excessive splitting

or from bringing to light previously unrecognized divergences via molecular work, but rather from exploration of new areas. It is important to note that “new areas” in this context should be taken to refer not only to remote geographic localities but also to unexplored biotic groups occurring in otherwise “well-studied” regions.

North American “microlepidoptera” represent the latter type of “new area.” Among insects, which are the most diverse eukaryotic group on earth, Lepidoptera is one of four enormously diverse orders, with approximately 10,000 described species in

(Hodges et al. 1983). Within the order, groups traditionally termed “”

(=Mimallonoidea through , sensu Kristensen et al. 2007) have been rather extensively documented. In contrast, Nearctic “microlepidoptera” (= Micropterigoidea through Thyridoidea, sensu Kristensen et al. 2007) remain poorly known. For example, a recent monograph (Hodges

1999) of Nearctic moths of the gelechiid Chionodes Hübner treated 187 species, of which

115 were described as new. Many other groups of microlepidoptera still await this basic, descriptive treatment.

Aside from mere documentation of species, knowledge of geographic range is vital to understanding organizational patterns of biodiversity, and such understanding is in turn critically important in applications such as identifying refugia and assessing human impact on particular species or natural systems (Donoghue and Alverson 2000). Basic information on geographic distribution of North American microlepidoptera is generally very incomplete. Many of the species listed by Hodges et al. (1983) are known from single localities and have not been recorded since the time of their original descriptions, this typically being 100 years ago or more.

“Rediscoveries” of such species often greatly enlarge previously known ranges (Metzler and

3 Zebold 1995), even to the degree of including new (Koster and Harrison 1997;

Heppner 1997; Landry 1998). It is therefore clear that expanding knowledge of geographic ranges of Nearctic microlepidoptera, and developing from that a useful understanding of

distributional patterns of these insects will not be served merely by databasing existing museum

specimens but will also require field investigation.

Any effort to establish a Lepidoptera knowledge base should emphasize microlepidoptera

not only because they are presently poorly known, but also because they represent an

overwhelmingly large percentage of the higher-level (suprageneric) diversity within the order.

Kristensen et al. (2007) grouped world Lepidoptera into 39 superfamilies, of which 31 are

microlepidoptera. For America north of Mexico, Hodges et al. (1983) listed 62 families of

Lepidoptera, of which 43 are microlepidopte. The latter also comprise 48% of the total species,

indicating that microlepidoptera make up a substantial proportion of Nearctic Lepidoptera

diversity even at lower taxonomic levels (and this does not take into account the relatively large

number of species that are as yet undescribed).

Microlepidoptera perform beneficial ecosystem services such as biological control

(Nuessly and Goeden 1984) and pollination (Reynolds et al. 2009; Yoder et al. 2010), and they

contribute to community diversity by serving as food for birds and other vertebrates (Baldwin

1970; Maher 1979; Wiens and Rotenberry 1979) and for many predaceous and parasitic

arthropod species (Okuyama 2007; Whitfield and Wagner 1991). Microlepidoptera also include

pests of economic importance, e.g., spruce budworm, Choristoneura fumiferana (Clemens),

diamondback moth, Plutella xylostella (Linnaeus), and pink bollworm, Pectinophora gossypiella

(Saunders).

4 That our basic knowledge of microlepidoptera remains so incomplete is particularly

unfortunate considering that many microlepidoptera groups have sizeable components in North

America, and among these are some, e.g., (Hsu and Powell 2005) and

(T. Harrison, unpublished data) in which the Nearctic fauna represents the majority of diversity worldwide. Likewise, in some groups (e. g., ), the basal taxa, which are of primary importance in phylogenetic analyses, are Nearctic (Davis 1987). Therefore, no accurate statement about any aspect of the entire order Lepidoptera will be possible until much additional work has been completed on North American microlepidoptera, and because of this critical importance, studies of the Nearctic component of these insects are as much needed as are those to be conducted in any other global region.

Of particular urgency are studies of specialized biotic communities of limited occurrence, especially those that now face the threat of serious degradation or even extinction. The unique informational perspectives offered by these systems, some of which were once the dominant communities in their respective regions, are vitally important to understanding the overall pattern and process of biodiversity well enough to manage it successfully.

Eastern tallgrass prairie in the Midwestern USA constitutes one such system. As recounted in Metzler et al. (2005), tallgrass prairie originated approximately 10,000 years ago in the Dakotas, Nebraska, , and (Delcourt and Delcourt 1980). With the advent of warmer, drier conditions, prairie began to extend eastward until, by 4000-5000 years ago, it had in some areas reached nearly to the east coast of the USA (Mehrhoff 1997). Finally, a westward retreat of prairie began about 3000-4000 years ago, this being brought about by a shift to the cooler, moister conditions that remain today (Winkler 1988). By the time of European settlement, tallgrass prairie had become restricted to an area encompassing southern

5 and Wisconsin, the northern two-thirds of Missouri, and most of , Illinois, and , with

substantial extensions into Michigan, , and . Today, an estimated 99% of original

prairie has been eliminated, mainly by conversion to agriculture (Swengel and Swengel 2001).

Because of the recognized importance of preserving the remaining tracts of prairie in the state,

much attention has been and continues to be focused on their conservation.

Hill prairies make up a designated subtype of eastern tallgrass prairie. They are island-

like openings on steep, otherwise forested slopes that face south or southwest (Evers 1955;

Kilburn and Warren 1963; Robertson et al. 1995). They occur along the River and

adjacent areas of tributaries, from Minnesota and Wisconsin southward to Iowa and southern

Illinois. In Illinois, they are found in scattered localities on the Mississippi River along the entire

western border of the state, along the Illinois River from north of Peoria southward to the

Mississippi River, and on the Sangamon River, a tributary of the Illinois (Evers 1955; Robertson

et al. 1995). A number of small hill prairies are also found on the Embarras River in east-central

Illinois (Vestal 1918; Reeves et al. 1978; Ebinger 1981).

In 1976-1977, the Illinois Natural Areas Inventory (INAI) project identified 446 hill

prairies within the state. Among these, 127 sites totaling 216.2 ha were found to be of high

quality (Grades A and B), these representing four types as defined by edaphic substrate: loess

(187.4 ha), glacial drift (20.8 ha), gravel (5.9 ha), and sand (2.1 ha) (Nÿboer 1981).

The vegetation of hill prairies can be characterized as a mixture of tallgrass prairie plants,

disjunctly occurring species that are otherwise found farther west in the plains region, and hill

prairie endemics (Schwartz et al. 2000). Lists of hill prairie plants were provided by Evers

(1955), Ebinger (1981), Robertson et al. (1995), and Schwartz et al. (2000). A three-way log-

linear model implemented by Schwartz et al. (2000) indicated that plant species composition of

6 Illinois hill prairies is similar to that of other eastern tallgrass prairie types (silt-loam, sand-

gravel), except that Bouteloua curtipendula () and Kuhnia eupatorioides (Asteraceae)

are common on hill prairies but not on other prairie types.

Hill prairies are of particular importance in conservation biology, because they represent

one of the most intact prairie systems remaining today. Robertson et al. (1995) speculated that,

because the steep slopes of hill prairies have prevented their being converted to row crops, the

percentage of original hill prairies remaining in Illinois is probably higher than that of other

prairie types. Furthermore, the spatial aspects of hill prairies are more similar to their original

form than are those of other prairie types, because hill prairies always existed as small, insular

fragments delimited by ravines on either side and by forest above and below (Schwartz et al.

2000).

Degradation of Illinois hill prairies is most often attributable to one of two causes.

Nÿboer (1981) found that, of 446 hill prairies surveyed for the INAI project during 1976-1977,

319 (71.5%) were severely degraded from livestock grazing. An additional threat to Illinois hill

prairie systems is invasion by woody plants (Kilburn and Warren 1963; White 1978; Voigt 1983;

Werner 1994). The rate of this type of degradation can be quite rapid; for example, McClain

(1983) calculated that, between 1937 and 1974, 62% of the collective area of five Jersey County

hill prairies had been converted to forest. Similarly, Schwartz et al. (2000) reported that between

1940 and 1988, large (39.2ha) and medium (1.7-72ha) hill prairie sites in Illinois averaged a 63%

decrease in size, whereas small sites (<0.5ha) averaged a 72% decrease. This decline agrees with

their observation that most hill prairies in the state became more fragmented during the study interval, and perimeter-to-area ratios showed an increase of over 100%, so that by 1988 the sites had become more vulnerable to woody invasion than they were 50 years earlier.

7 For approximately the past 30 years, a focus of discussion and research in conservation

biology has been the so-called SLOSS (single large or several small) debate. This debate stems

from the application to conservation biology of the theory of island biogeography (MacArthur

and Wilson 1963) in determining optimal design and management strategies (generally taken to

be those that will allow maximum number of species per area) for insular remnants of habitat,

such as hill prairies. No consensus has been reached in this debate, and it appears that optimal

reserve design must take many factors into account. Cutler (1994) found that a large, species- rich reserve would conserve more species than any combination of smaller reserves, but only of the archipelago were perfectly nested (that is, if the species component of a small fragment is a subset of that of the next larger fragment), whereas, if the structure shows deviation from perfect nestedness, a large reserve might have no advantage over several smaller ones. Baz and Garcia-

Boyero (1996), on the other hand, determined that species richness of in oak forest fragments in Spain would best be conserved by creating many small, scattered reserves.

Kunin (1997) tested the effects of reserve shape on plant distribution and concluded that reserves should become more elongate with increased size, rather than being circular as had been previously suggested. Virolainen et al. (1998), in a study of vascular plants in boreal mires, emphasized the importance in habitat assessment of considering rarity and taxonomic diversity, in addition to the traditionally used criterion of species richness. Haddad (1999a,b, 2000) and

Haddad and Baum (1999) observed higher densities of butterflies in habitats connected by corridors than in isolated fragments.

Small hill prairies are evidently as important as large sites in capturing and conserving biodiversity. Species-area relationships observed by Schwartz et al. (2000) indicated that plant diversity in very small hill prairies (<0.5ha) is proportionately similar to that seen in larger ones.

8 They also found that small sites contain higher numbers of infrequent plant species than expected

[Kilomogorov-Smirnov test of deviations from expected frequencies generated using the methodology of Simberloff and Gotelli (1983)], and a chi-square test of association of common species and site indicated that common species are likewise present, as expected, on small sites.

Management of prairie remnants has been the subject of considerable discussion, and in recent years, research has been undertaken to determine the effects of various management practices, particularly the use of prescribed burning, on prairie . Metzler et al. (2005) provided a list of literature on the use of fire in prairie management. The results of this research have been quite varied. For example, Swengel (1999) found that haying and grazing are more favorable to prairie-specialist butterflies than is burning. Other studies, however, have documented rapid recolonization by arthropods even after burns that caused nearly 100 percent mortality (Tooker and Hanks 2004) or have found that fire is not permanently harmful to arthropod populations if adjacent patches are left unburned to allow recolonization (Harper et al.

2000). Still other studies have shown actively beneficial effects of fire (Hartley et al. 2007;

Rooney 2010). The emergent picture is that, because of varying responses to fire by different prairie types, and by different species, a burning regimen must be tailored to the particular management goals that are in place for each reserve.

In this study, I surveyed and compared Pyraloidea (=Pyralidae + Crambidae sensu

Munroe and Solis 1998) and Tortricidae in two disjunct hill prairies in western Illinois. Previous lists of proposed prairie-restricted microlepidoptera have been dominated by these groups. For example, 37 of 39 microlepidoptera species that were suggested as prairie specialists by Reed

(1996) are “Pyralidae” or Tortricidae. Likewise, Metzler’s (2005) list covers 43

microlepidoptera species, of which 35 belong to these groups. Other groups of microlepidoptera

9 (e.g., some families of ) may harbor substantial numbers of prairie-associated

species as well, but information on the prairie components of these groups (e.g., the description

in the present study of two previously unrecognized leadplant-feeding species of Gelechiidae,

Chapters 4 and 5) is only beginning to come to light.

The majority of Nearctic species of Pyraloidea and Tortricidae for which life histories are

known feed as larvae on plants in only a single family (Forbes 1923; Robinson et al. 2010).

Given this, the present study is intended to explore two primary questions: (1) Do the two prairies support similar species diversities of Pyraloidea and Tortricidae, or does the larger, more floristically diverse prairie (Revis) harbor a higher diversity of these moths than does the smaller prairie with fewer plant species (Pere Marquette); and (2) Do Pyraloidea and Tortricidae display low beta diversity (i.e., similar sets of species) between the two sites, or do different species complements of these moths occur in the two respective prairies?

Materials and Methods

Microlepidoptera were collected at two loess hill prairies, Revis and Pere Marquette, occurring approximately 240 km apart, both lying along a northeast-to southwest line that is coincident with the course of the Illinois River. Both prairies were rated as being high-quality by the INAI. The sites also had equivalent management histories, in that neither had been burned, grazed, or hayed for more than five years prior to the collection dates. The composition of grass species was similar, and leadplant, Pursh (Fabaceae), was abundant at both sites. Diversity and abundance of Asteraceae, however, were high at Revis, low at Pere

Marquette, and slimflower scurfpea, Psoralidium tenuiflorum (Pursh) Rydberg (Fabaceae), was

10 common at Revis, absent from Pere Marquette, whereas the reverse was true for roundheaded bush clover, Lespedeza capitata Michx. (Fabaceae). The prairies also differed in size and latitude. Revis prairie, in Mason County, T20N, R7W, Sections 25, 26, and 36, is a 95.1 ha hill prairie. Pere Marquette prairie, in Jersey County, T6N, R11W, Section 9, is a 21.9 ha site. As to the surrounding vegetaitonal matrix, Revis is bordered by deciduous forest on one side, agricultural land on the other, whereas Pere Marquette is completely surrounded by a large, continuous expanse of deciduous forest.

A very limited amount of rearing of larval moths was done in this study, as time allowed. Larvae were reared in 40 x 45mm plastic cups that were placed inside sealed plastic bags to preserve humidity levels. Collections of adult moths were made on seven dates (6 June,

14 June, 29 June, 7 July, 2 August, 10 August, and 1 September, 2005) using an 18.9-liter bucket trap equipped with an 8-watt UV light (BioQuip Products, Rancho Dominguez, CA). One trap was placed approximately in the center of each prairie. On each trapping date, both prairies were sampled on the same night. Collections were made on seven nights, from early June through early September. Although only Tortricidae and Pyraloidea were identified for analysis in the present study, all microlepidoptera were removed and pinned from each sample. Moths were identified by sight or with the aid of genital dissection. Genital preparations were stored in glycerin in 96-well immunoassay trays (Becton Dickinson Labware, Franklin Lakes, NJ).

Voucher specimens will be deposited into the collection of the Illinois Natural History Survey,

Champaign, IL. Holotypes and other primary type material of species that are described as new will be deposited into the collection of the National Museum of Natural History,

Washington, DC.

11 Information on microlepidoptera collected in this survey was compiled as a file in

Microsoft Office Excel 2003. Alpha diversity of moths was calculated using the Shannon-

Wiener index, H’ = Σ Pi * LN(Pi), where Pi = proportion of moth community in the ith species

category. Sorenson’s index was used to determine beta diversity: C = (2 x jN) / (aN + bN),

where aN, bN, and jN are number of species in community a, community b, and communities a and b, respectively. Values of C can range from 0 (maximum beta diversity, in which no species

are shared between a and b) to 1.0 (minimum beta diversity; all species shared between a and b).

Analysis of variance was used to test for differences in moth diversity and density among

collection dates.

Results

A total of 2798 moths representing 214 species of Pyraloidea and Tortricidae were

collected. Numbers by collection date (Table 1) generally reflect the trend usually seen in

microlepidoptera flight period in central Illinois, with a peak in June and early July, followed by

a lull in late July and August, and a moderate resurgence in early September (Godfrey et al.

1987). Alpha and beta diversity values are presented in Table 2. Alpha diversity is similar in the

two sites, both for total combined sample and for Pyraloidea or Tortricidae alone. Beta diversity

values, however, indicate low species complementarity between the two sites, especially for

Tortricidae. Tortricidae collected at Revis are predominated by species associated with prairie

forbs (Fabaceae and Asteraceae), whereas, with the exception of the leadplant-feeding

Hystrichophora taleana, tree-feeding species are most prevalent among Tortricidae collected at

Pere Marquette.

12 Among Pyraloidea, the leadplant-feeding phycitine, Sciota rubescentella (Hulst)

occurred at both sites. Similarly, three species of Peoria (P. approximella (Walker), P. bipartitella Ragonot, and P. gemmatella (Hulst)), all of which feed on grasses as larvae, were collected in both prairies. The Monarda-feeding laticlavia (Grote and Robinson) was collected only at Revis, as were Australephestiodes stctella (Hampson), Eurythmia angulella

(Zeller), Phycitodes reliquella (Dyar), and Argyria nummulalis Hübner. Life histories are unknown for the latter four species, but they are assessed here as likely candidates for being prairie-dependent species because of legume-feeding (or, in the case of A. nummulalis, grass-

feeding) habits of species that are held to be taxonomically related (Heinrich 1956; Martinez and

Brown 2007). The absence of these species from Pere Marquette might indicate that their larval

host plants occur only at Revis, but in the case of A. nummulalis, size of the prairie might be a

determining factor; observations by T. Harrison indicate that A. nummulalis generally is seen

only in large prairies.

Among Tortricidae (Table 3), the samples from Revis were dominated by prairie-

associated Asteraceae- and Fabaceae-feeding species (the latter being the two species of

Hystrichophora) that reflect the high diversity of forbs of these families at Revis. The fact that

Hystrichophora vestaliella (Zeller), the life history of which is unknown, is present in abundance

at Revis but absent from Pere Marquette agrees with a pattern observed by T. Harrison, i. e., that

this moth is seen only on prairies where Psoralidium tenuiflorum occurs. I therefore propose that

P. tenuiflorum is the most likely candidate for being the larval host plant of H. vestaliana on hill

prairies in Illinois. At Pere Marquette, the commonest tortricids were tree-feeding species (Table

3). This undoubtedly reflects the fact that Pere Marquette is completely surrounded by a large

and diverse expanse of deciduous forest, with the light trap being in relatively close proximity to

13 the larval hostplants of the tree-feeding tortricids than at Revis because of the smaller size of

Pere Marquette prairie. An exception, however, is the leadplant-feeding Hystrichophora taleana

(Grote), which follows the same pattern as the leadplant-feeding Anacampsis (Gelechiidae) that is described elsewhere in this study, in that it was collected in greater numbers at Pere Marquette than at Revis. This might be simply a concentration effect, but even if so, it invites research to determine whether it is a result of differences in dispersion of egg deposition, or of distance of attraction to light.

Data from each of the seven collection dates were compared by analysis of variance to test for the effect of date on moth abundance per species (Table 4). No significant differences were seen, either in all data combined, Revis + Pere Marquette by taxon, Pyraloidea +

Tortricidae by site, or either Pyraloidea or Tortricidae by site. Typically, in moth samples collected at light, most species are collected in small numbers, with a few species being abundant

(Thomas and Thomas 1994; Thomas 1996; Summerville and Crist 2008). As a result, mean numbers of individuals per species tend to be similar among samples, regardless of absolute number of species per sample. This, in conjunction with the fact that the presence of a few abundant species amidst a matrix of many uncommon species generates high standard deviations, appears to be the reason that no significant differences were observed.

Discussion

Even though the two surveyed prairies are separated by only 240 km north-south distance, latitude might in some cases be a factor in determining microlepidoptera species composition. One such case is the leadplant-feeding Filatima sp. (Gelechiidae) that is described

14 as new elsewhere in this study. Despite the fact that leadplant is a dominant species at both sites, the Filatima is abundant at Revis Prairie but absent from Pere Marquette. A second undescribed

species of Filatima is known from far-northern Illinois, where it is associated with Hudsonia sp.

(Cistaceae) (J. R. Wiker, in litt.), and yet another undescribed Filatima has been reared from

silky prairie clover, Dalea villosa (Nutt.) Spreng. in Minnesota (M. J. Hatfield and J. van der

Linden, in litt.). This could suggest a tendency toward northern geographical ranges for some

Filatima spp. in the Midwest, in which case Revis prairie might represent the southern extremity

of distribution for the leadplant-feeding species. Testing this hypothesis lies beyond the scope of

this study, but I note that surveying leadplant for Filatima larvae on prairies north and south of

Revis in early to mid-June would be an easy and straightforward matter, due to the conspicuous larval feeding damage and unmistakable, boldly-striped of this moth.

In conclusion, I recognize this study is limited, in that only two prairies were compared. However, this preliminary examination of the microlepidopteran component of two previously unsurveyed sites allows a number of observations and recommendations. First, an operational problem in this study was that, because, despite the hardware-cloth screen that was placed over the UV light bulb to exclude larger insects, the samples very often contained enormous numbers of smaller insects, especially certain species of Coleoptera and Trichoptera, which damaged the moths. As a result, an inordinately high percentage of microlepidoptera specimens retrieved from light traps were in such poor condition as to require dissection in order to be identified. One solution to this problem would be to install the necessary personnel to collect moths manually at each site on each collecting date (Axmacher and Fiedler 2004). Even if this were logistically and economically possible, it probably would introduce considerable variation in expertise in recognizing and capturing the moths. An approach that might

15 realistically solve the problem is to collect moths in standard light traps, and then DNA barcode

the moths (Hebert et al. 2003; Janzen et al. 2005) before any attempt is made to identify them via

dissection. In concept, with this a priori sorting, only one individual of each species would need

to be dissected for identification. Barcoding might also confer the advantage of shedding light

on cryptic species complexes that involve species that are collected on prairies, e.g., Epiblema

strenuana (Walker) and albiguttana (Zeller) (Gilligan et al. 2008).

Second, the dearth of life-history information on moths collected in this survey

recommends that an effort should be made to engage in a dedicated program of rearing prairie moths, either from collecting larvae on the prairies, from obtaining ova from adult female moths, or from caging known or suspected larval food plants. This is a very open area of exploration.

For example, in my study, rearing was done incidentally to light trapping, yet even this casual

rearing effort yielded two previously undescribed species of Gelechiidae that are obligately

associated with tallgrass prairie. Such information, once acquired and disseminated,

immeasurably increases the value of light-trap data for the species in question.

Third, as was shown by Schwartz et al. (2000) for plants, my results indicate that even

small tracts with comparatively low plant diversity should not be disregarded, in that they can

serve as refugia for some species of prairie-dependent microlepidoptera.

Fourth, controlled, replicated experimental work is needed, to determine which

conditions, above and beyond mere plant composition, are important in maintaining

microlepidoptera diversity in prairies, so that, if possible, these conditions can be taken into

account during the process of charting management strategies.

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Table 1. Counts of moths (number of species, followed by number of individuals) by collection date, for Pyraloidea and Tortricidae collected at Revis and Pere Marquette hill prairies.

Revis Pere Marquette Pyraloidea Tortricidae Total Pyraloidea Tortricidae Total 6VI 25, 251 21, 244 46, 495 30, 151 13, 78 43, 229 14VI 14, 22 16, 126 30, 148 33, 283 12, 38 45, 321 29VI 30, 241 26, 73 56, 314 8, 35 4, 25 12, 60 7VII 18, 64 17, 111 35, 175 13, 26 10, 31 23, 57 2VIII 14, 127 7, 202 21, 329 13, 92 5, 33 18, 125 10VIII 11, 79 6, 23 17, 102 15, 67 5, 23 20, 90 1IX 21, 126 15, 110 36, 236 18, 88 14, 29 32, 117 Total 75, 910 71, 889 146, 1799 75, 742 40, 257 115, 999

24 Table 2. Alpha and beta diversity values for Pyraloidea and Tortricidae collected at Revis and Pere Marquette hill prairies. Alpha Pyraloidea Tortricidae Total Revis 3.93 3.44 4.38 Pere Marquette 3.82 3.57 4.32 Beta 0.41 0.25 0.34

Table 3. Prevalent species of Tortricidae collected in light traps at Revis and Pere Marquette prairies. Species Number of individuals Revis Hystrichophora vestaliana (Zeller) 124 Pelochrista pallidipalpana (Kearfott) 124 Eucosma matutina (Grote) 15 Pelochrista corosana (Walsingham) 36 Eucosma albiguttana (Zeller) 22 Eucosma giganteana (Riley) 7 Hystrichophora taleana (Grote) 7 Eucosma ridingsana (Robinson) 5 Pere Marquette Hystrichophora taleana (Grote) 59 Cydia caryana (Fitch) 37 Argyrotaenia quercifoliana (Fitch) 25 Cydia latiferreana (Walsingham) 19 Gymnandrosoma punctidiscanum Dyar 18

Table 4. Results (p values) of analysis of variance to test for effect of collection date on moth abundance per species. Site All moths Pyraloidea Tortricidae Both sites 0.235 0.845 0.109 Revis 0.141 0.454 0.099 Pere Marquette 0.735 0.817 0.564

25 Chapter 2. Evaluation of Prairie-Dependent Microlepidoptera of the Basal Families

Micropterigidae through Gelechiidae

Abstract. Basal microlepidoptera have not conventionally been a focus of Lepidoptera research on prairies despite the fact that they comprise a very large number of species. Based upon both published information and my preliminary sampling of microlepidoptera on prairies in

Illinois, I provide a general evaluation for each of these groups, as to the likelihood that it contains prairie-dependent species. Of the 31 families considered, 12 are evaluated as possible, and two as likely or certain, to include prairie-dependent species.

Key words. Microlepidoptera, sub-Apoditrysia, prairie dependent-species, larval host plants,

Coleophoridae, Gelechiidae

Introduction

Basal microlepidoptera of the families Micropterigidae through Gelechiidae are frequently overlooked in biotic surveys, because their small size and concealed feeding habits make them difficult to collect and identify. Over 2500 described species of these moths occur in

North America (Hodges et al. 1983), and some groups that never have been comprehensively monographed are known to harbor large numbers of undescribed species. For example, based on examination of museum specimens, Landry (1991) estimated the actual number of species of

Scythridinae (Xylorictidae) to be 400 to 500, as opposed to 35 species listed by Hodges (1983).

Given their impressive species diversity, moths of these groups undoubtedly play key roles in many food webs. Feeding specialization and limited dispersal capacity may place them at

26 particular risk in endangered habitats such as prairies. Here I evaluate the likelihood of prairie

endemicity among basal microlepidoptera.

No asterisk = unlikely; * = possible; ** = probable or certain.

Micropterigidae.: The larvae feed on lichens in deciduous forest (Davis 1987a). Two species of Micropterigidae are native to North America; only one of these, Epimartyria auricrinella occurs in the eastern USA. It has not been recorded from Illinois but has been collected in Indiana, only a few kilometers from the eastern border of Illinois. It probably occurs in the northern and eastern parts of Illinois, especially in unglaciated sites that harbor liverwort

beds in heavily shaded mesic areas at the bases of sandstone cliffs.

Eriocraniidae: Larvae are leaf miners on trees (Davis 1978). One species, Dyseriocrania

griseocapitella, is common and widespread in Illinois. The larva is a full-depth blotch

on black oak, Quercus velutina (Fagaceae). Smaller trees (under six meters in height) are

preferred, and most leaf mines occur at a height of about two meters above ground level. The

leaf mine is very similar to the mines of some chrysomelid , in that it is rather puffy and

contains a good deal of stringy frass. The moth is univoltine. In central Illinois, the larva matures in early May. It then leaves the mine and burrows underground, where it spins a small ovate cocoon, inside which it remains as a larva until the following spring, when it pupates and then emerges as an adult in late April.

Hepialidae: The larvae are root borers in trees, or moss feeders (Wagner 1987).

Hepialidae are absent or rare in Illinois.

27 /: larvae are leaf or bark miners on woody plants (Braun 1917;

Wilkinson 1979; Wilkinson and Scoble 1979; Wilkinson 1981; Wilkinson and Newton 1981;

Newton and Wilkinson 1982; Davis and Stonis 2007). Adults are usually darkly colored, often with one or more white or metallic fasciae on the forewing. Nepticulids of some species are among the smallest Lepidoptera known

Incurvariidae//: These moths occur primarily in the far-western USA

(Davis 1967; Powell 1969; Davis et al. 1992); the few species that occur in the East are associated with deciduous forest. Larvae for which life histories are known are internal feeders, often in the reproductive tissue of the host plant.

Heliozelidae: leaf miners on woody plants (Davis 1987b).

*Tischeriidae: Larvae feed as leaf miners, mainly on , Fagaceae, or Asteraceae

(Braun 1972; Puplesis and Diškus 2003). The family is well represented in Illinois, and the

Asteraceae-feeding habit could indicate prairie-dependent species, but there is no evidence of this at present.

*: Larvae of most species of the large genus Acrolophus for which life histories are known are reported to feed on roots of grass (Hasbrouck 1964), but the genus is mostly western, and only a few species are known from Illinois.

*: Larvae of the relatively few species for which life histories are known tend to feed on dead-wood or on animal products (Robinson 2009). However, one species,

Eccritothrix trimaculella (Chambers), appeared in some numbers in light traps in this study.

*: Larvae are most species are leaf miners; those of the genus Caloptilia are leaf rollers. Larvae feed on many plant species, with a preponderance toward woody plants

(Braun 1908; Forbes 1923). Acrocercops

28 pnosmodiella (Busck) (Fig. 1A) feeds on Onosmodium spp. (). In the present study, uhlerella (Fitch) (Fig. 1B) was collected in light traps, and leaf mines assumed to be those of this species were found on leadplant, Amorpha canescens Pursh

(Fabaceae). Porphyrosela desmodiella (Clemens) (Fig. 1C) has been reared from Lespedeza capitata Michx. (Fabaceae) in prairie (T. Harrison, unpublished data) but also from other, non- prairie legumes (Braun 1908).

* (Fig. 1D): Larvae are internal feeders (stem borers or leaf miners) in a wide variety of forbs and woody plants (Braun 1963); a large segment of Bucculatrix species feed on plants in the family Asteraceae. Adults were occasional in light traps in this study.

Metzler et al. (2005) included Bucculatrix simulans Braun and B. staintonella Chambers as possibly being prairie-associated species. The former feeds on common sunflower Helianthus anuus L. (Asteraceae), the latter on poplars, Populus spp. (Salicaceae).

Superfamily : The larvae of Attevidae, Yponomeutidae, ,

Acrolepiidae, , and Argyresthiidae feed mostly on woody plants. Larvae in forb- feeding groups (e.g., , , Heliodinidae) feed primarily on non-prairie

Hydrophyllaceae and Boraginaceae, , and Nyctaginaceae, respectively (Gaedike

1983, 1990; Heppner 1985, 1987; Hsu and Powell 2005; Landry 2007).

Superfamily Gelechioidea:

* (sensu Clark 1941, Hodges 1974): Depressariinae could have prairie species on or Asteraceae; Metzler et al. (2005) included Agonopterix pergandeella

(Busck) (life history and habitat unknown) as possibly being a prairie-associated species. Adults of this group were depauperate in light traps in this study.

29 *Ethmiidae (sensu Powell 1973) (Fig. 1E): Many species feed externally on leaves of

Hydrophyllaceae or Boraginaceae, but species occurring in the eastern USA tend to be forest species. However, one species, Ethmia longimaculella (Chambers), feeds on puccoons,

Lithospermum spp. (Boraginaceae) in open sand areas. Several species of Ethmia are known from Illinois, but no adults were collected in this study.

* (sensu Braun 1948; Kaila 1995a,b, 1996, 1997, 1999): Larvae of almost all species are leaf miners on grasses or sedges, which might suggest the presence of species associated with tallgrass prairie, but no Elachistidae were collected in this study.

* (Hodges 1978; Koster 2010): (Fig. 1F) includes many grass- and sedge-feeding species; adults of several species of Cosmopterix, including C. minutella Beutenmüller (life history unknown) were collected in light traps, occasionally in considerable numbers.

**Coleophoridae: RESEARCH WARRANTED

Subfamily Coleophorinae (Heinrich 1923): This is a large group centered around the genus Coleophora (145 described species in the Hodges et al. 1983 checklist). Larvae are internal feeders (mostly borers or leaf miners) in a wide array of both forbs and woody plants. I observed the larva of a Coleophora sp. (possibly undescribed) mining leaves of slimflower scurfpea, Psoralidium tenuiflorum (Pursh) Rydberg (Fabaceae) at a hill prairie in Mason County, and I found a fairly high diversity of adults in light traps in this study.

Subfamily Blastobasinae: life histories are unknown for most species, but some are associated with grasses (Adamski and Brown 1989). Adults appearing to represent several different species were collected in this study.

30 Subfamily Momphinae (Stehr 1987) (Fig. 1G) are stem, flower, or fruit borers, or leaf miners, with a proclivity for feeding on Cistaceae and Onagraceae. Examination of museum specimens by T. Harrison (unpublished data) indicates that there are many undescribed species.

In this study, adults of two undescribed species of Momphinae were collected in hill prairies in

Mason and Jersey Counties, respectively.

*Glyphidoceridae: Almost no life-history information is available for moths of this small group. Metzler et al. (2005) included Glyphidocera wrightorum Adamski and Metzler as being possibly a prairie-associated species.

*: Nothing is known of the life histories of most species in this small group. Metzler et al. (2005) included inquisitor Hodges as possibly being a prairie- associated species. T. Harrison (unpublished data) collected a related species, Chedra pensor

Hodges, in an open sand prairie in Mason County, Illinois. Life histories are unknown for both of these moths.

*Xylorictidae (Subfamily Scythridinae) (Fig. 1H): Life histories are unknown for most species, but some are known to feed as leaf miners or skeletonizers on Asteraceae (Landry

1991). Scythridines were rare in light-trap samples in this study, but this would be expected due to the diurnal habits of adults in this group.

**Gelechiidae: RESEARCH WARRANTED

Larvae of the majority of species in this large family (886 described species in North

America, according to the Lee et al. 2009 checklist) feed on leaves as tiers, rollers, or miners, but many other modes of feeding are seen (Busck 1903; Hodges 1986, 1999). Metzler et al. (2005) included elegantella (Chambers) and Gnorimoschema huffmanellum Metzler and

Adamski (life histories unknown) as possibly being prairie-associated species. Danderson et al.

31 (2007) investigated plant-herbivore interactions of Coleotechnites eryngiella Bottimer (Fig. 1I),

which feeds in flowers of rattlesnake master, Eryngium yuccifolium Michaux (Apiaceae) in Illinois.

An undescribed species of Monochroa, and the seldom-collected Sophronia primella Busck (Fig.

1J), were collected in numbers in this study. Life histories of these species are unknown, but some Sophronia spp. in other world regions feed on grasses (Robinson et al. 2010). Two prairie- restricted leadplant feeding spp. are described as new in the present study (Chapters 4 and 5).

Additional unrecognized prairie species probably are present; a high diversity of adults was seen in the light traps.

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Figure 1. Representatives of some basal microlepidoptera families in which prairie-dependent

species might be included (photos not to same scale). A, Acrocercops pnosmodiella

(Gracillariidae); B, Macrosaccus uhlerella (Gracillariidae); C, Porphyrosela desmodiella

(Gracillariidae); D, Bucculatrix eugrapha (Bucculatricidae); E, Ethmia zelleriella (Ethmiidae sensu Powell 1973); F, Cosmopterix clemensella (Cosmopterigidae); G, Mompha passerella

(Coleophoridae); H, Scythris basilaris (Xylorictidae); I, Coleotechnites eryngiella (Gelechiidae);

J, Sophronia primella (Gelechiidae).

38

39 Chapter 3. Moth Diversity in Three Biofuel Crops and Native Prairie

Abstract. The expanding demand for biofuel feedstock may lead to large-scale conscription of land for monoculture production of biofuel crops with concomitant substantial negative impacts on biodiversity. I compared moth diversity in light-trap samples from corn, miscanthus, switchgrass, and native prairie, to determine whether there is an observable relationship between plant species diversity and moth abundance and diversity. Moth alpha diversity was highest in prairie and was higher in switchgrass than in the other two biofuel crops.

Beta diversity generally was low among the biofuel crops, and prairie shared lower beta diversity

with switchgrass than with corn or miscanthus. Analysis of variance showed no significant

differences in moth abundance per species among treatments. The alpha and beta diversity-index

findings are consistent with those of other studies on arthropods in biofuel crops and provide

evidence to suggest that large-scale conversion of acreage to biofuel crops may have substantial

negative effects on arthropod biodiversity both within the cropping systems and in the

surrounding landscape.

Key words. Bioenergy, biodiversity, moths, agricultural landscapes, ecosystem services

Introduction

In recent years, a dramatic increase in the price of oil has spurred great interest in developing alternative liquid fuels based on plant biomass. Under the directives of the US

Energy Independence and Security Act of 2007, production of renewable fuels will increase from

30 million cubic meters in 2008 to 135 million cubic meters in 2022, with an estimated

40 100,000,000 ha of US land being engaged in this production (Perlack et al. 2005; Graham et al.

2007; Schmer et al. 2008).

Such large-scale conscription of land for monoculture production of biofuel crops inevitably will have a substantial negative impact on biodiversity (Cook et al. 1991; Naylor et al.

2007; Tilman et al. 2009). As summarized by Landis and Werling (2010), alteration of habitat

(especially reduction of plant species diversity) by biofuel cropping systems can cause food web shifts (Hedgren 2007) and changes in arthropod community structure (Nitterus and Gunnarsson

2006; Ulyshen and Hanula 2009). This can result in increased impact of arthropods, whether they be existing pests (Mattson et al 2001; Coyle 2002; Hanson 2003) or species that newly emerge as pests (Dimou et al. 2007; Ward et al. 2007). As well, the efficacy of beneficial arthropods in limiting pest numbers may be reduced, in that biofuel crops alter their spatial or temporal distribution (Frank et al. 2008) or fail to provide them with needed shelter or alternative food sources (Carmona et al. 1999; Menalled et al. 2001; Gardiner et al. 2010).

Moreover, the effects of large-scale monocultures on arthropod biodiversity can extend beyond crop plantings, into the surrounding landscape. Pests may build up in number and spill over into other crops (Ahmad et al. 1984; Semere and Slater 2007), and the lack of untreated refuge areas can intensify selection pressure on pest populations to develop insecticide resistance

(Hansen 2003). Concomitantly, a reduction in arthropod-mediated ecosystem services such as pollination and pest control can be seen in the surrounding landscape (Redderson 2001; Landis et al. 2008), and negative effects on organisms of other trophic levels (e.g., birds that depend on arthropods for food) also have been observed (Sage and Tucker 1997, 1998).

In terms of number of species, insects comprise the largest group of organisms on earth, with over 1 million species described, and probably an even greater number as yet

41 undescribed (Anonymous 2011). Order Lepidoptera (butterflies and moths) is one of the four

largest groups of insects. Recent tallies of the number of described Lepidoptera species

worldwide range from 146,277 (Heppner 1991) to 160,000 (Kristensen et al. 2007, who

estimated the actual total at 500,000). The most recent check list of North American Lepidoptera

(Hodges et al. 1983) tabulated 11,233 species, or 13 percent of described Nearctic insect species

(Kosztarab and Schaefer 1990). Moths make up the overwhelming majority (93 percent) of

Lepidoptera species in North America.

Because of their enormous species diversity, and the great abundance of some species,

moths occupy an important role in many different community interactions. For example, larval

and/or adult moths are a substantial dietary component of wasps (Whitfield and

Wagner 1991), spiders (Okuyama 2007), grassland birds (Baldwin 1970; Maher 1979; Wiens

and Rotenberry 1979), rodents (Nuessly and Goedart 1984), and bats (Dai et al. 2009). Adult

moths can be important as pollinators, especially in specific associations with certain plant

species (Reynolds et al. 2009; Yoder et al. 2010). It also has been postulated that feeding

damage inflicted by moth larvae serves to prevent particular plant species from becoming overly

dominant within their respective communities (Metzler et al. 2003).

Moths are singularly dependent upon the plants that occur within a community. Larvae

of most moth species feed obligately on living plant tissue, and a large proportion of these

phytophagous species display some degree of hostplant specificity (Dyer et al. 2007). Therefore,

a positive relationship between plant diversity and moth diversity is likely to exist. To test this

hypothesis, I compared diversity of adult moths in native prairie vegetation and in three biofuel

crops: corn (Zea mays L.), miscanthus (Miscanthus x giganteus), and switchgrass (Panicum virgatum L.). My expectation was that prairie vegetation, with its higher plant species diversity,

42 harbors a greater diversity and abundance of moths than do the crop plantings. This study is the

first to examine impacts of non-corn biofuel feedstock production on moth diversity.

Materials and Methods

Sites from which moths were collected are listed in Table 5. At the University of

Illinois site south of Savoy (2007 collections), plantings occur in 0.9 ha plots of switchgrass,

miscanthus, mixed miscanthus and vetch, and corn, with six plots per treatment. At the

University of Illinois Energy Biosciences Institute Energy Farm (2008 and 2009 collections),

switchgrass, miscanthus, corn, and mixed prairie are planted in alternating plots of 0.69 ha each,

with four plots per treatment. Planting began at the Savoy plots in 2004, at the EBI Energy Farm

in 2008. At the Agricultural Centers (2009 collections), miscanthus and switchgrass are planted

in eight adjacent, alternating subplots of 100m2 each. These plots were planted in 2002 at

DeKalb and Dixon Springs, and in 2004 at Fairfield, Brownstown, and Orr. At all of these sites,

the miscanthus and switchgrass plots are adjacent to corn in large-scale production (40+ ha).

Each of the matched prairie remnants for Brownstown, Orr, and DeKalb is approximately 40 km from the Ag Center. Two of these prairies, Byler Cemetery and Temperance Hill Cemetery, are small (approximately 0.9 ha in area). Twelve-Mile Prairie, on the other hand, occupies a large area (92 ha) overall, but it occurs as a long (ca. 19 km, as the name implies), very narrow strip

(ca. 30 m in width) between Illinois Route 37 and the track of the Illinois Central Railroad. All three of these sites, as well as the reconstructed prairie plots at the University of Illinois Energy

Farm, support plant species that are characteristic of mesic prairie (Moorehouse and Hassen

43 2004). Handel (1996) provided a list of dominant prairie plants that occur in the mesic prairie

community (Table 6).

Collections were made using an 18.9-liter bucket trap equipped with an 8-watt UV light

(BioQuip Products, Rancho Dominguez, CA). One trap was placed approximately in the center

of each plot; at each site, one plot per vegetation system was sampled, on the same night. In

2007, moths were sampled on ten nights, from 12 June to 7 September; in 2008, two collections

were made, on 24 July and 20 August; and in 2009, each site was sampled on one night, the dates

ranging from 24 June through 19 August. Moths were identified by sight or with the aid of genital dissection. Genital preparations were stored in glycerin in 96-well immunoassay trays

(Becton Dickinson Labware, Franklin Lakes, NJ). Voucher specimens will be deposited into the collection of the Illinois Natural History Survey, Champaign, Illinois.

Alpha diversity of moths was calculated using the Shannon-Wiener index, H’ = Σ Pi *

LN(Pi), where Pi = proportion of moth community in the ith species category. Sorenson’s index

was used to determine beta diversity: C = (2 x jN) / (aN + bN), where aN, bN, and jN are number

of species in community a, community b, and communities a and b, respectively. Values of C

can range from 0 (maximum beta diversity, in which no species are shared between a and b) to

1.0 (minimum beta diversity; all species shared between a and b).

Results

A total of 5411 moths, representing 252 species in 25 families, were

collected over the course of this study (Table 7). Alpha diversity values are presented in Tables

8 and 9, beta diversity values in Table 10. In the 2007 collections, alpha diversity values are

44 very similar among all crops, and beta diversities are low (almost 70 percent species complementarity in all pairings). In 2008, alpha diversity is highest for prairie and lowest for miscanthus and corn, with switchgrass intermediate between them. Beta diversity is lowest in prairie x switchgrass and miscanthus x switchgrass (56 and 57 percent species complementarity, respectively) and highest in corn x miscanthus (29 percent). In 2009, alpha diversity is higher in prairie and switchgrass than in miscanthus and corn. The Shannon-Wiener value of 3.25 for the combined prairie data and 2.12 for the 19 August prairie data, despite the fact that the highest number of moth species and individuals occurred in prairie, is due to the fact that one species,

Stereomita andropogonis (Gelechiidae), makes up 42 percent (549 individuals) of the combined sample. All of these individuals of S. andropogonis were collected at Twelve-Mile Prairie on 19

August. When S. andropogonis is removed from the combined 2009 prairie data, the Shannon-

Wiener value is 4.41, and when removed from the prairie data for 19 August, the value is 3.77.

Beta diversity values are similar for all pairings (48-56 percent complementarity), with corn x prairie slightly higher (41 percent).

Data from each of the collection dates were compared by analysis of variance to test for the effect of treatment on moth abundance per species (Table 11). No significant differences were observed, either in all data combined, Revis + Pere Marquette by taxon, Pyraloidea +

Tortricidae by site, or either Pyraloidea or Tortricidae by site. Usually, in light-trap samples of moths, a few species are abundant, whereas the majority of species are collected in small numbers (Thomas and Thomas 1994; Thomas 1996; Summerville and Crist 2008). Because of this, regardless of absolute number of species per sample, mean numbers of individuals per species generally are similar among samples. Also, the presence of a few abundant species

45 amidst a matrix of many uncommon species generates high standard deviations; this, along with similar among-sample means, appears to be the reason that no significant differences were seen.

Discussion

The recent, increased demand for corn as a biofuel feedstock (Westcott 2007) is bringing about marked changes to agricultural landscapes in the Midwestern USA. In the interest of maximizing production, growers are planting corn not only on former soybean acreage, but also on land that formerly was left fallow or used in the production of various minor crops (Landis et al. 2008). Concerns have arisen that the resulting reduction in diversity and abundance of native plants in and among agricultural tracts might have an adverse effect on arthropods and the ecological services that they impart, in agricultural systems and beyond. As a result of these concerns, research has begun to emerge to examine the degree of impact that might be expected.

Studies to date have focused on the effect of agricultural landscape on beneficial arthropods, especially pollinators, and predaceous and parasitic species that have the potential to serve as biological control agents of agricultural pests. In general, these studies have borne out the hypothesis that agricultural landscapes that offer a diverse flora of native perennials will harbor a higher diversity of beneficial arthropods than will extensive monocultures of annual plants such as corn and soybean. For example, a metaanalysis by Bianchi et al. (2006) found that simple agricultural landscapes have fewer natural enemies and greater pest pressure (76 percent and 45 percent of studies, respectively) than do diverse agricultural landscapes that offer relatively large areas of noncrop habitat, and a field survey by Gardiner et al. (2010) found that

46 beneficial arthropods were more diverse and abundant in native prairie and in switchgrass than in

corn.

Some of my results appear to diverge from the patterns seen in previous studies, but in

large part, such apparent anomalies can be explained on basis of methodological conditions. For

example, the extremely similar alpha-diversity values and high species complementarities in the

2007 sample data are a reflection of the small size and immediate proximity of the plots, and of

the relatively extensive sampling effort (ten light-trapping dates). In the 2008 and 2009 samples,

the somewhat elevated species complementarity between switchgrass and miscanthus (57 and 56

percent, respectively) is also probably the result of sampling from small, proximate plantings of

these two crops. Finally, the high beta diversity of corn x miscanthus in 2008 (29 percent

complementarity) likely reflects the fact that these two samples have the lowest species counts

(17 species each), so that the effect of each species difference is magnified, compared to the

situation in a more species-rich sample.

Overall, the data from my moth collections are consistent with previous studies, in that, in both of the years (2008, 2009) in which prairie vegetation was compared with the three biofuel crops, the ranking of moth diversity and abundance was prairie>switchgrass>corn/miscanthus.

Plant species diversity likely is an important factor in my results. Planting corn in straight, uniformly-spaced rows readily allows active control of weeds, so that a mature cornfield presents a closed canopy of uniform height, overshadowing a large expanse of bare soil. Mixed prairie, switchgrass, and miscanthus are established either by using a drill or by broadcast seeding. This results in a dense dispersion of plants, with bare soil representing only about five percent of the total area (B. Robertson, unpublished data, cited in Gardiner et al. 2010). Miscanthus, like corn, forms a closed canopy at a uniform height of over 2m, under which plant diversity is low.

47 Prairie and switchgrass, on the other hand, present a very heterogeneous canopy height, from

only a few cm to over 2m, and plant diversity is high compared to that of corn or miscanthus.

This height differential might suggest that my results are an artifact of differential visibility of

the light traps in the various crops, but my data do not support that hypothesis, because the 2008

sampling at the EBI Energy Farm was done when the plots were in the initial stages of

establishment, such that light-trap visibility from edge of plot was equivalently high in prairie,

switchgrass, and miscanthus, and low in corn. If trap visibility were an influential factor, then

lower diversity and abundance of moths should have been in corn than in miscanthus, but such

was not the case (17 species, 85 individuals were collected in both crops). Similarly, lower

diversity should have been found in miscanthus in 2009 (crop established at full height) than in

2008, but, again, this was not seen (17 species, 85 individuals in 2008, versus 31 species, 150

individuals in 2009).

Continued research is needed to determine how best to manage agricultural landscapes in ways that cost-effectively deliver maximal benefits of arthropod-mediated ecosystem services

(Landis et al. 2000; Gurr et al. 2004; Samways 2007; Whittingham 2007; Isaacs et al. 2009). An excellent review of this complex problem is provided by Landis and Werling (2010).

Furthermore, it is vitally important to keep growers apprised of all available information coming from this research, because they are most likely to adopt conservation-oriented management practices if they are able to make informed decisions on how to proceed in a way that will confer the greatest possible economic benefit (Gurr et al. 2003; Olson and Wäckers 2007; Isaacs et al.

2009). Finally, as pointed out by other workers (Fiedler et al. 2008; Isaacs et al. 2009), studies such as this one, which demonstrate benefits of conservation management practices to native biodiversity above and beyond beneficial arthropods, can be valuable in promoting acceptance,

48 advocacy, and implementation of these practices among non-grower landowners and society at large.

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Table 5. Sites from which moths were collected (counties in parentheses). University of Illinois plots were sampled in 2007, EBI Energy Farm 2008-9, all others 2009.

University of Illinois plots (south of Savoy) Miscanthus, corn, and switchgrass, on same site; no prairie

EBI Energy Farm (south of Urbana) Crop grasses and reconstructed prairie on same site

Orr Agricultural Research and Demonstration Center (Pike) Matched prairie remnant: Byler Cemetery Prairie (Adams)

Northern Illinois Agronomy Research Center (DeKalb) Matched prairie remnant: Temperance Hill Cemetery Prairie (Lee)

Brownstown Agronomy Research Center (Fayette) Matched prairie remnant: Twelve-Mile Prairie at Farina (Fayette)

University of Illinois Extension Office at Fairfield (Wayne) Matched prairie remnant: None

Dixon Springs Agricultural Center (Johnson) Matched prairie remnant: None

56 Table 6. Dominant prairie plant species characteristic of mesic prairie community, from which moths were sampled in this study (from Handel 1996).

Family Species Asteraceae Silphium laciniatum Helianthus grosseserratus Liatris pycnostachya Prenanthes aspera Solidago juncea Poaceae Rudbeckia hirta Andropogon gerardii Elymus virginicus Panicum virgatum Sorghastrum nutans Fabaceae Baptisia lactea Desmodium sessiflifolium Desmodium illinonese Apiaceae Eryngium yuccifolium Monarda fistulosa Gentianaceae Gentiana puberulenta Euphorbiaceae Euphorbia corollata Onagraceae Gaura longifolia

Table 7. List of moths collected in the study.

Family Subfamily Genus Species Opostegidae Pseudopostega quadristrigella Tischeriidae Astrotischeria sp. 1 Acrolophidae Acrolophus popeanellus Acrolophidae Acrolophus mortipennellus Acrolophidae Acrolophus texanellus Acrolophidae Amydria onagella Tineidae Homostinea curviliniella Psychidae Thyridopteryx ephemeraeformis Gracillariidae Acrocercops astericola Gracillariidae Caloptilia violacella Gracillariidae Leucospilapteryx venustella Gracillariidae Parectopa plantaginisella Gracillariidae Parornix sp. 1 Bucculatricidae Bucculatrix agnella Bucculatricidae Bucculatrix angustata Bedelliidae somnulentella Attevidae Atteva pustulella

57 Plutellidae Plutella xylostella Glyphipterigidae Diploschizia impigritella Oecophoridae Epicallima argenticinctella Elachistidae Antaeotricha schlaegeri Elachistidae Antaeotricha leucillana Elachistidae Elachista sp. 1 Elachistidae Gonioterma mistrella Amphisbatidae Psilocorsis reflexella Glyphidoceridae Glyphidocera lithodoxa Glyphidoceridae Glyphidocera septentrionella Coleophoridae sp. 1 Coleophoridae Blastobasis sp. 2 Coleophoridae Coleophora mayrella Coleophoridae Coleophora sp. 1 Coleophoridae Coleophora sp. 2 Coleophoridae Coleophora sp. 3 Coleophoridae Coleophora sp. 4 Coleophoridae Coleophora sp. 5 Coleophoridae Hypatopa sp. 1 Coleophoridae Mompha circumscriptella Coleophoridae Mompha murtfeldtella Coleophoridae Pigritia sp. 1 Xylorictidae Scythris trivinctella Cosmopterigidae Cosmopterix minutella Cosmopterigidae Cosmopterix pulcherrima Cosmopterigidae phragmitella Cosmopterigidae concolorella Cosmopterigidae miscecolorella Cosmopterigidae sexnotella Gelechiidae Anacampsis fullonella “roseosuffusella” Gelechiidae Aristotelia 1 “roseosuffusella” Gelechiidae Aristotelia 2 Gelechiidae Chionodes discoocellella Gelechiidae Gelechiidae Chionodes sp. 1 Gelechiidae Chionodes sp. 2 Gelechiidae Chionodes sp. 3 Gelechiidae Dichomeris ligulella Gelechiidae Dichomeris isa Gelechiidae Dichomeris juncidella Gelechiidae Dichomeris inversella Gelechiidae Frumenta nundinella Gelechiidae Helcystogramma hystricella Gelechiidae Gnorimoscheminae sp. 1

58 Gelechiidae Gnorimoscheminae sp. 2 Gelechiidae Gnorimoscheminae sp. 3 Gelechiidae Isophrictis sp. 1 Gelechiidae Isophrictis sp. 2 Gelechiidae Isophrictis sp. 3 Gelechiidae Monochroa pullusella Gelechiidae Monochroa sp. 1 Gelechiidae Phthorimaea operculella Gelechiidae Polyhymno luteostrigella Gelechiidae Sinoe robiniella Gelechiidae Stegasta bosqueella Gelechiidae Stereomita andropogonis Gelechiidae Syncopacma palpilineella Gelechiidae sp. 1 Gelechiidae sp. 2 Tortricidae Ancylis metamelana Tortricidae Ancylis sp. 1 Tortricidae Argyrotaenia velutinana Tortricidae Argyrotaenia quercifoliana Tortricidae Bactra furfurana Tortricidae Bactra verutana Tortricidae Carolella bimaculana Tortricidae Tortricidae Choristoneura rosaceana Tortricidae Clepsis persicana Tortricidae Clepsis peritana Tortricidae Conchylis oenotherana Tortricidae Endothenia hebesana Tortricidae Epiblema sp. 1 Tortricidae Epiblema sp. 2 Tortricidae Epiblema sp. 3 Tortricidae Epiblema sp. 4 Tortricidae Epiblema sp. 5 Tortricidae Epiblema sp. 6 Tortricidae Episimus argutanus Tortricidae Eucosma vagana Tortricidae Eucosma cataclystiana Tortricidae Eucosma agricolana Tortricidae Eucosma glomerana Tortricidae Eucosma derelicta Tortricidae Eucosma fiskeana Tortricidae Eucosma sombreana Tortricidae Eucosma albiguttana Tortricidae Eucosma matutina Tortricidae Tortricidae Lobesia carduiella

59 Tortricidae Tortricidae Pelochrista pallidipalpana Tortricidae Phaneta parmatana Tortricidae Phaneta raracana Tortricidae Phaneta ochrocephala Tortricidae Phaneta ochroterminana Tortricidae Phaneta ornatula Tortricidae Phaneta striatana Tortricidae Platynota idaeusalis Tortricidae Platynota flavedana Tortricidae Sparganothis sulfureana Tortricidae Xenotemna pallorana Tortricidae Cochylinae sp. 1 Tortricidae Cochylinae sp. 2 Tortricidae Cochylinae sp. 3 Tortricidae Cochylinae sp. 4 Tortricidae Cochylinae sp. 5 Tortricidae Cochylinae sp. 6 Tortricidae Cochylinae sp. 7 Tortricidae Cochylinae sp. 8 Tortricidae Cochylinae sp. 9 Tortricidae Cochylinae sp. 10 Tortricidae Cochylinae sp. 11 Tortricidae Cochylinae sp. 12 Tortricidae Cochylinae sp. 13 Oidaematophorus sp. 1 Pterophoridae Pyraloidea Achyra rantalis Pyraloidea Aethiophysa lentiflualis Pyraloidea Chrysoteuchia topiarius Pyraloidea Condylolomia participialis Pyraloidea Crambus agitatellus Pyraloidea Crocidophora tuberculalis Pyraloidea Donacaula sp. 1 Pyraloidea Elophila gyralis Pyraloidea Eoreuma densella Pyraloidea Euchromius ocelleus Pyraloidea Eurythmia hospitella Pyraloidea Fissicrambis hemiochrellus Pyraloidea Fissicrambis mutabilis Pyraloidea Goya stictella Pyraloidea Homeosoma ellectellum Pyraloidea Hymenia perspectalis Pyraloidea Hypsopygia costalis Pyraloidea Pyraloidea Mimoschinia rufofascialis

60 Pyraloidea Neodactria luteolellus grp. 1 Pyraloidea Neodactria luteolellus grp. 2 Pyraloidea Nomophila nearctica Pyraloidea Oenobotys vinotinctalis Pyraloidea Ostrinia nubilalis Pyraloidea Parapediasia teterrella Pyraloidea Pyraloidea Peoria bipartitella Pyraloidea Peoria gemmatella Pyraloidea Petrophila sp. 1 Pyraloidea Phycitodes mucidella Pyraloidea Platytes vobisne Pyraloidea Pococera sp. 1 Pyraloidea Pyrausta signatalis Pyraloidea Pyrausta tyralis Pyraloidea Pyrausta orphisalis Pyraloidea Scoparia basalis Pyraloidea Sitochroa sp. 1 Pyraloidea Synclita obliteralis Pyraloidea Synclita sp. 2 Pyraloidea Synclita sp. 3 Pyraloidea Udea rubigalis Pyraloidea Urola nivalis Pyraloidea sp. 1 Pyraloidea sp. 2 Geometridae Anavitrinella pampinaria Geometridae Chlorochlamys chloroleucaria Geometridae Eupithecia miserulata Geometridae Haematopis grataria Geometridae Idaea demissaria Geometridae Orthonama obstipita Geometridae Orthonama centrostrigaria Geometridae Geometridae Semiothisa multilineata Geometridae Synchlora aerata Geometridae sp. 1 Arctiidae Apantesis vittata Arctiidae Crambidia sp. Arctiidae Cisseps fulvicollis Arctiidae Estigmene acrea Arctiidae Holomelina sp. Arctiidae Phragmatobia lineata Arctiidae Pyrrharctia isabella Arctiidae Spilosoma virginiana Agrotis ipsilon Noctuidae Amolita fessa

61 Noctuidae Noctuidae Noctuidae Caenurgina erechtea Noctuidae Noctuidae Crambodes talidiformis Noctuidae Elaphria festivoides Noctuidae Faronta rubripennis Noctuidae Feltia subgothica Noctuidae Galgula partita Noctuidae Helicoverpa zea Noctuidae Heliothis virescens Noctuidae Noctuidae Hormoschista latipalpis Noctuidae Noctuidae Idia aemula Noctuidae Lacinipolia renigera Noctuidae Leucania phragmatidicola Noctuidae Leucania multilineata Noctuidae Leucania scirpicola Noctuidae Lithacodia carneola Noctuidae Noctuidae Noctuidae Melanchra picta Noctuidae Noctuidae Noctuidae Orthodes crenulata Noctuidae Noctuidae Peridroma saucia Noctuidae Phalaenostola larentioides Noctuidae Plathypena scabra Noctuidae Platysenta vecors Noctuidae Noctuidae Pseudaletia unipuncta Noctuidae Pseudoplusia includens Noctuidae Renia sp. 1 Noctuidae arcigera Noctuidae Schinia lynx Noctuidae Schinia thoreaui Noctuidae Simyra henrici Noctuidae Spodoptera ornithogallii Noctuidae Spodoptera frugiperda Noctuidae Spragueia apicalis Noctuidae Noctuidae Tarachidia candefacta Noctuidae Tetanolita floridana Noctuidae

62 Noctuidae Thioptera nigrofimbria Noctuidae Zanclognatha protumnusalis Noctuidae sp. 1 Noctuidae sp. 2 Noctuidae sp. 3 Noctuidae sp. 4

Table 8. Alpha diversity values for moths collected in biofuel crops on each sampling date of the study. Zero indicates that the site was sampled but no moths were collected; “---“ indicates no sample, either because there was no matched prairie or (miscanthus 23 July 2009) because of equipment failure. See note in the results section regarding the value for the 19 August 2009 sample.

Date Prairie Switchgrass Miscanthus Corn 2007 --- 12 June --- 2.91 2.92 2.86 20 June --- 2.55 2.37 2.84 29 June --- 2.42 2.19 2.52 5 July --- 2.14 2.96 1.55 30 July --- 1.16 1.57 1.16 10 August --- 2.65 2.95 2.04 15 August --- 3.13 3.49 3.06 24 August --- 3.46 2.91 3.16 30 August --- 1.79 1.74 1.49 7 September --- 2.93 2.64 2.87 2008 24 July 2.20 2.04 1.41 0.00 24 August 2.72 1.90 1.48 1.71 2009 24 June 3.32 2.94 2.80 2.82 23 July --- 2.87 --- 1.79 31 July --- 2.91 1.39 2.35 5 August 2.97 0 0 0.87 13 August 3.93 3.03 2.26 1.61 19 August 2.12 3.31 3.39 3.25

63 Table 9. Shannon-Wiener values for EBI light-trapped moths, with data combined by year. Numbers in parentheses indicate number of species, followed by number of individuals. See note in Results section regarding the value for the 2009 prairie sample.

2007 2008 2009 Prairie ------(48, 345) 2.93 (155, 1314) 3.25 Switchgrass (120, 788) 3.98 (25, 153) 2.18 (110, 524) 4.16 Miscanthus (123, 894) 3.96 (17, 85) 1.68 (75, 284) 3.71 Corn (115, 689) 3.69 (17, 85) 1.74 (70, 233) 3.70

Table 10. Sorenson’s index values for EBI light-trapped moths. PG = prairie; SW = switchgrass; MXG = miscanthus; MZ = corn.

2007 2008 2009 PG SW MXG PG SW MXG PG SW MXG SW --- 0.56 0.52 MXG --- 0.69 0.41 0.57 0.48 0.56 MZ --- 0.67 0.66 0.41 0.43 0.29 0.41 0.53 0.56

Table 11. Results of analysis of variance to test for effect of collection date on moth abundance per species. 2007 2008 2009 Date p value Date p value Date p value 12 June 0.936 23 July 0.769 24 June 0.973 20 June 0.555 19 August 0.607 23 July 0.207 29 June 0.961 31 July 0.408 5 July 0.408 5 August 0.657 30 July 0.876 13 August 0.240 10 August 0.865 19 August 0.469 15 August 0.780 24 August 0.517 30 August 0.615 7 September 0.739

64 Chapter 4. A New, Prairie-Restricted Species of Filatima Busck (Lepidoptera:

Gelechiidae) from Illinois

Abstract. Filatima revisensis (Gelechiidae) is described from individuals collected as larvae feeding inside shelters constructed of silked-together leaflets of leadplant, Amorpha canescens

(Fabaceae). Filatima revisensis is bivoltine; overwintering occurs in the larval stage. Because this insect is restricted to tallgrass prairie, it is likely to be of concern to conservation biologists.

In the interest of naming this moth and clarifying its identity, a description is provided, and diagnoses are given to differentiate it from F. ornatifimbriella, F. xanthuris, F. adamsi, and F. occidua, all of which are externally similar to F. revisensis.

Key words. Amorpha canescens, microlepidoptera, Gelechioidea, taxonomy, North America, tallgrass praire, habitat-restricted species, conservation biology

Introduction

The genus Filatima Busck (Lepidoptera: Gelechiidae) is a large, primarily Holarctic assemblage, the majority of Nearctic species occurring in semiarid regions of the western USA and Mexico

(Hodges and Adamski 1997). Hodges (1983) listed 76 species of Filatima for North America north of Mexico. This number is an underestimate, as no comprehensive taxonomic revision has been published for the Nearctic component of the genus. Because some Nearctic Filatima species have yet to be described, and because biologies are known for relatively few species, accurate identification of moths belonging to this genus can be problematic. Adding to the

65 problem of identification is the fact that Filatima includes groups of externally similar species, these groups not necessarily being monophyletic.

Hodges and Adamski (1997) provided redescriptions of two externally similar Filatima species,

F. ornatifimbriella (Clemens) and F. xanthuris (Meyrick), along with original descriptions of two additional species, F. adamsi and F. occidua, that are similar in appearance to F. ornatifimbriella and F. xanthuris. Based on genital morphology, Hodges and Adamski concluded tentatively (pending a complete phylogenetic analysis of Filatima) that these four species do not represent a monophyletic unit. In the context of baseline taxonomy and identification as in the present paper, however, it is useful to consider this assemblage of species together. It is referred to here as the ornatifimbriella color group.

During the course of my study of microlepidoptera of tallgrass prairies, I reared an undescribed

Filatima species of the ornatifimbriella color group from larvae feeding on leadplant, Amorpha canescens Pursh (Fabaceae), a prairie-restricted plant. The insect appears to be monophagous, in that larvae have been observed to feed neither on other prairie legumes nor on non-prairie species that are taxonomically proximate to leadplant, e.g., false wild indigo,

(L.) (which is the larval host plant of F. ornatifimbriella). Through its obligate association with leadplant, the undescribed moth is restricted to a biotic community, tallgrass prairie, that is of interest to conservation biologists because of its present extremely limited occurrence. It is advisable, therefore, that the leadplant-feeding Filatima should be named, to facilitate communication regarding the moth, and that diagnoses should be provided to differentiate it from the other four species of the ornatifimbriella color group, some of which are known or

66 likely to be sympatric with it (Hodges and Adamski 1997). The purpose of this paper is to provide a description and diagnosis of the leadplant-feeding Filatima species.

Materials and Methods

Larvae were reared individually in one-ounce size (40 x 45mm) snap-lid plastic cups enclosed in one-quart size (17 x 20 cm) self-sealing plastic bags. Each adult moth was pinned on a minuten pin, spread, and double-mounted on a foam staging block, with its associated larval and pupal exuviae preserved in a gelatin capsule affixed to the main mounting pin. Genitalic dissection of adult moths followed the procedure of Clarke (1941). In addition, to allow maximal visualization with minimal distortion of the male genitalia, the ventral part (valvae and vinculum) of the genitalia was separated from the tegumen at one of its connection points with that structure, after which it was flattened for study and illustration as per the recommendations of Pitkin (1984). I separated the left valve at its connection with the tegumen, in order to preserve the left-right chirality of the asymmetrical valve/vinculum assembly. This greatly facilitates discussion of these structures, compared to a dissection in which the separation is made at the right valve, as was done by Hodges and Adamski (1997), in which case the chirality of the asymmetrical structures is reversed and potentially confusing (e.g., Hodges and Adamski

(p. 42) state that the saccus of F. adamsi is directed toward the right, as it appears in their Fig.

18, whereas it actually is directed toward the left). That my method presents the tegumen and associated structures in dorsal aspect, with left-right chirality reversed, is not problematic, because, in a cleared preparation of Filatima, all of the structures in question are equally visible in dorsal and in ventral aspect, and also because this part of the genitalia is symmetrical, such

67 that chirality never is a point of discussion. Photographs were taken using a Canon EOS Digital

Rebel camera attached to a Bausch and Lomb dissecting microscope, or an Olympus compound microscope. Wing measurements were done on a personal computer in Photoshop CS (Adobe,

San Jose, CA), by comparing photographs of wings and a millimeter ruler that were taken at a standard uniform magnification under a dissecting microscope.

Results

Filatima revisensis Harrison, new species

(Figs. 2-6)

Diagnosis. The three species of the ornatifimbriella color group for which life histories are known can be differentiated by their respective larval host plants (all Fabaceae): Amorpha canescens (F. revisensis), Amorpha fruticosa (F. ornatifimbriella), and Thermopsis, Lupinus,

Robinia, and Vicia (F. xanthuris). Male adult F. revisensis is readily differentiated from other species of the ornatifimbriella color group (except F. occidua, the male of which is unknown) by the presence of three separate projections (single costal lobe, plus two projections of saccular lobe) on each valve. In F. ornatifimbriella, F. xanthuris, and F. adamsi, one or both valvae bear only two projections (single costal and saccular lobes). Female F. revisensis is differentiated from other species of the ornatifimbriella color group (except F. adamsi, the female of which is unknown) by the presence of a signum (signum absent in F. xanthuris), presence of an invaginated pocket near each of the anterior apophyses (pockets absent in F. occidua), and the large, complex signum (signum a small, simple crescent-shaped sclerite in F. ornatifimbriella).

68 Additionally, adult F. revisensis and F. occidua differ in their respective geographic distributions

(Midwestern USA, versus Washington and California, USA), coloration (brown, versus gray), and size (FW length 6.1-8.0 mm, versus 5.5-6.2 mm), and F. revisensis and F. adamsi differ in

their respective geographic distributions (Midwestern USA, versus Maine, USA)

Description. Adult (Fig. 2). Head. Ocellus present; each scale of vertex and frons pale brown

basally, dark brown apically (vestiture of such scales referred to hereafter as "bicolored brown");

haustellum brown basally, then paler brown for most of its length; maxillary palpus brown; labial

palpus bicolored brown, mixed with pale-ochreous scales, the latter predominating on

ventromedial surface; second segment with "furrowed brush" scale tuft as typical for genus.

Thorax. Dorsum and tegulae bicolored brown; ventral surface shining pale ochreous with a few

brown scales. Legs. Lateral surface gray intermixed with pale ochreous, tibia with pale-

ochreous band at midlength and apex, medial surface shining pale ochreous, all tarsomeres

except fifth with pale-ochreous ring at apex; hind tarsus margined dorsally from near base with a

row of long yellowish hairlike scales, inception of row marked by a pale-ochreous blotch on

dorsolateral surface of tibia. Forewing. Mean length, 7.1mm (range, 6.1-8.0 mm, n=6); dorsal

surface uniformly bicolored brown, in some individuals with rather extensive suffusion of gray;

four dark-brown spots, located as follows: single spot (faint in some individuals) immediately

posterior to fold at 0.25 wing length; two spots, separated by fold, at 0.40 wing length, posterior

spot located slightly more basally than anterior one; single spot midway between anterior and

posterior margins of wing at 0.60 wing length; costal margin gray interspersed with pale-

ochreous scales, apical margin to tornus gray but lacking pale-ochreous scales; fringe gray, basal tier of scales dark tipped, forming a line; ventral surface uniformly shining grayish brown.

69 Hindwing. Dorsally and ventrally, wing shining pale brown, not appreciably darkened apically; fringe shining pale brown. Abdomen. Dorsally, anterior three visible segments (actual segments

2-4) yellow, remaining segments brown with shining-gray posterior margins, color change between yellow and brown segments coincident with a change in scale morphology (Fig. 3); venter whitish, heavily intermixed with brown scales. Male eighth abdominal segment modified

(Fig. 4), the sternite 0.7x as long as wide, with short anterolateral arms, posterior margin rounded and shallowly and narrowly emarginate medially, the tergite 1.6x as long as wide, with elongate anterolateral arms, posterior margin rounded to a broad medial point. Male genitalia (Fig. 5).

Uncus and gnathos typical for the genus, the former hoodlike, the latter short, narrow, and hooklike, its posterior margin serrate; vinculum asymmetrical, saccus directed toward left; costal lobes of valvae similar, each lightly sclerotized, elongate and narrow, with slightly-dilated apex clothed ventrally with a brush of recurved setae; saccular lobes asymmetrical, each divided into an elongate, narrow anteroventral projection and a short, broad medial projection; anteroventral and medial projections of left valve heavily sclerotized, the former longer and slightly broader than anteroventral projection of right valve, the latter nozzle shaped, shorter and broader than medial projection of right valve, and with subapical thornlike development on posterior margin; aedeagus sclerotized and bulbous at base, the sclerotized region narrowing posteriorly into a strip that extends along left side of aedeagus to its posterior extremity, where the strip is abruptly recurved anteroventrad; a T-shaped sclerite immediately ventrad of the sclerotized strip, a large midventral thornlike sclerite immediately posterad of the T-shaped sclerite, and a small thornlike sclerite posterodorsad of the midventral sclerite, near apex of aedeagus. Female genitalia (Fig.

6). Ovipositor elongate (length 11.5 x width), posterior apophyses 3.6x as long as anterior apophyses, membrane in region of anterior apophyses invaginated into a pair of deeply-

70 invaginated pockets bearing microtrichia; ductus bursae sclerotized, with many microtrichia,

reflected asymmetrically to left, narrowing to midlength then dilating to reach broadest point at

anterior end; corpus bursae and accessory bursae membranous; signum large, appearing as

curved medial band, divided into anterior and posterior regions, the former more heavily

sclerotized than the latter, the band with a pair of inwardly-directed, triangular lateral lobes.

Larva. Length of mature larva approximately 11mm. Coloration typical of larvae of this genus: head and prothoracic shield orange, body whitish with paired dorsal, subdorsal, and lateral blackish stripes.

Biology. Larva on Amorpha canescens. A row of opposing leaflets (often an entire leaf) is silked together to form a feeding chamber; late-stage feeding damage is characterized by browning of the affected leaflets. Pupation occurring either inside the larval shelter or (usually) outside the shelter in a frass-covered silken cocoon. Bivoltine, overwintering as mature larva.

Pupation of overwintered larvae occurs in central Illinois probably in early May, as first- generation adults are seen in mid- to late May. Second-generation larvae mature and pupate in mid-June; adults emerge, mate, and oviposit in early July; second-generation larvae, which will overwinter, mature in early to mid-August.

Distribution. Presently known from tallgrass prairies in Mason, Menard, and Morgan Counties,

Illinois, USA.

Types. Holotype male. Collected as larva on Amorpha canescens, USA, Illinois, Mason

County, Revis Hill Prairie, 15-VI-2005, T. Harrison, emerged 3-VII-2005; [red label]

HOLOTYPE/Filatima revisensis ♂/Harrison. Allotype female. Same data as for holotype,

71 except emerged 9-VII-2005; [yellow label] ALLOTYPE/Filatima revisensis ♀/Harrison.

Paratypes. Three males, same data as for holotype, emerged 2-VII-2005, 3-VII-2005 and 4-VII-

2005; [each with blue label] PARATYPE/Filatima revisensis ♂/Harrison. All types deposited into the insect collection of the United States National Museum of Natural History.

Etymology. The species is named for Revis Hill Prairie, Mason County, Illinois, the site from which it originally was reared.

Discussion

Because no phylogenetic analysis of Filatima has been undertaken, character-state polarities have not been assigned; thus, species relationships within the ornatifimbriella color group are difficult to assess. In this group, different morphological characters align with different sets of species. For example, the form of the signum of F. revisensis is similar to that of F. occidua, whereas evaginated pockets near the anterior apophyses are shared by F. revisensis, F. ornatifimbriella and F. xanthuris but are absent in F. occidua. Taxonomic proximity of larval host plants (in the species for which their identity is known) similarly does not parallel morphological similarity, as the two Amorpha feeders, F. ornatifimbriella and F. revisensis, are not seen to share more character states than either shares with other species of the group. In fact, some character states in F. revisensis (e.g., development of an elongate, narrow anteroventral projection of the saccular lobe of each valve) are unique within the group. Compounding the problem is the fact that two species of the group, F. adamsi and F. occidua, are known from individuals of only one gender. Therefore, given the rather bewildering array and/or absence of

72 morphological information that is presented when only the species of the ornatifimbriella color

group are examined, I concur with Hodges and Adamski (1997) that "until a phylogenetic

analysis is completed for all Filatima, a suitable hypothesis of relationships among species

cannot be made."

Literature Cited

Clarke, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera.

Bulletin of the Brooklyn Entomological Society 36: 149-161.

Hodges, R. W. 1983. Family Gelechiidae. Pp. 19-25 in Hodges, R. W., T. Dominick, D. R.

Davis, D. C. Ferguson, J. G. Franclemont, E. G. Munroe, and J. A. Powell, eds. 1983.

Check List of the Lepidoptera of America North of Mexico Including Greenland. London: E.

W. Classey Limited and The Wedge Entomological Research Foundation, 284 pp.

Hodges, R. W., and D. Adamski. 1997. The identity of Filatima ornatifimbriella

(Clemens 1864) (Gelechioidea: Gelechiidae). Journal of the Lepidopterists' Society

51: 32-46.

Klots, A. B. 1970. Lepidoptera. Pp. 115-130 in Tuxen, S. L., ed. Taxonomist's

Glossary of Genitalia in Insects, 2nd Edition. Copehagen: Munksgaard, 359 pp.

Pitkin, L. 1984. A technique for preparing complex male genitalia in microlepidoptera.

Entomologist's Gazette 37: 173-179.

Figures

Figure 2. Filatima revisensis. Adult, dorsal aspect. Scale bar = 5.0mm.

73

Figure 3. Filatima revisensis. Scale morphology of adult abdominal terga. A, overview showing difference in coloration in segments 2-4 versus the posterior segments; B and C, detail of scales occurring on 2nd through 4th terga; D, detail of scales occurring on posterior terga.

Figure 4. Filatima revisensis. Sclerites of eighth abdominal segment of male adult. A, sternite, ventral aspect, flattened; B, tergite, dorsal aspect, flattened.

Figure 5. Filatima revisensis. Male genitalia, dissected to present valvae and vinculum in lateral and ventral aspects, respectively, tegumen and associated structures in dorsal aspect, aedeagus removed; CL, costal lobe of valve; MP, medial projection of saccular lobe of valve; AP, anteroventral projection of saccular lobe; VI, vinculum; UN, uncus; GN, gnathos; TE, tegumen;

AE, aedeagus, ventral aspect.

Figure 6. Filatima revisensis. Female genitalia, ventral aspect. OV, ovipositor; PA, posterior apophysis; DB, ductus bursae; AA, anterior apophysis; AB, accessory bursae; SI, signum; CB, corpus bursae.

74

75

76

77 Chapter 5. A New, Prairie-Restricted Species of Anacampsis Curtis (Lepidoptera:

Gelechiidae) from Illinois

Abstract. Anacampsis wikeri (Lepidoptera: Gelechiidae), new species, is described. The larva of A. wikeri feeds on leaves of a prairie legume, Amorpha canescens (Fabaceae). The moth is univoltine, with mature larvae occurring in late May, adults in June; overwintering occurs in the adult stage. The adult of A. wikeri is externally very similar to that of another prairie- associated legume-feeding species, Anacampsis psoraliella; sight identification of adults of these two species that have not been reared thus is rendered problematic. Larval host plant specificity and adult genital morphology, however, allow unequivocal diagnosis. These characters are discussed, and male and female genitalia are illustrated, for both species.

Key words. Microlepidoptera, taxonomy, larval hostplants, Fabaceae, Amorpha, Psoralea,

Pediomelum, Psoralidium, Orbexilum, prairie ecology, conservation biology

Introduction

The genus Anacampsis Curtis (Lepidoptera: Gelechiidae) is primarily Holarctic in distribution.

It is well represented in America north of Mexico, where 23 described species are known to occur (Lee et al. 2009). Larvae of the majority of species in the genus are monophagous or oligophagous; for example, nine out of 13 Nearctic Anacampsis species listed by Robinson et al.

(2009) feed on plants of only one respective genus.

78 Dependence upon one or a few related plant species by a phytophagous insect can result in

obligate association of the insect with a particular biotic community. Some communities occur

as tracts that are so restricted in number and size that they have become the subject of active

efforts aimed at their preservation and management. During the course of my study of microlepidoptera of one such community (tallgrass prairie), Illinois lepidopterist James R. Wiker

brought to my attention a moth that he reared from a larva feeding on leaves of leadplant,

Amorpha canescens Pursh (Fabaceae), a prairie-restricted legume. The moth proved to be an

undescribed species of Anacampsis. The adult of the leadplant-feeding Anacampsis is externally very similar to that of a described species, Anacampsis psoraliella Barnes and Busck (Fig. 7), which similarly feeds on prairie-associated Fabaceae (namely, several species formerly ascribed to the genus Psoralea).

In developing an effective management strategy, it is advisable to be aware, to the greatest possible degree, of the ecological profiles of all biotic components that occur in the community in question. Therefore, to bring this insect to the attention of biologists who are involved with studies and/or management of prairie communities, to delineate the life history of the moth, to provide information that will allow its accurate identification, and to name the species so that

communication regarding it can be facilitated, I provide a description of the leadplant-feeding

Anacampsis. For convenience of comparison and diagnosis, I also discuss and illustrate the male

and female genitalia of A. psoraliella.

79 Materials and Methods

Larvae were reared individually in 40 x 45 mm snap-lid plastic cups, these being sealed inside 17

x 20 cm self-sealing plastic bags. Adult moths were spread and double mounted on foam staging

blocks, with their associated larval and pupal exuviae preserved in a gelatin capsule affixed to

the main pin. Terminology regarding genital morphology follows Klots (1970). For genitalic dissection, I used the procedure that was described by Clarke (1941). Also, the ventral part

(valvae and vinculum) of the genitalia was separated from the tegumen and flattened for study

and illustration as recommended by Pitkin (1984). Genital preparations were stored in glycerin

in genitalia vials affixed to the main pin of the associated specimen. For photography, a Canon

EOS Digital Rebel camera was attached to a Bausch and Lomb dissecting microscope or to an

Olympus compound microscope. Wing measurements were done in Photoshop CS (Adobe, San

Jose, CA), by comparing photographs of wings with a millimeter ruler that was photographed at

the same magnification as the wing. Scientific names of plants were obtained from the PLANTS

database (USDA, NRCS 2009).

Results

The adult of A. psoraliella differs structurally from that of the leadplant-feeding species. In the

male genitalia (Fig. 9), the valve is relatively broad and is abruptly narrowed near the apex; the

juxta is laterally constricted near its anterior end and is quadrate in shape, with the posterior

margin entire; the vinculum is relatively broad and U-shaped; and the aedeagus in lateral aspect

is relatively narrower along its entire length. In the female genitalia (Fig. 11), a conical

80 sclerotized invagination is absent at the ostium bursae; the ventral surface of A8 bears a curved, longitudinally-oriented sclerite on either side of the midline; the tergal projection of A8 is straight and dorsoventrally flattened along its entire length; and a small, bandlike signum is present on the corpus bursae.

Anacampsis psoraliella also differs from the leadplant-feeding species in larval foodplant preference. So far as is known, the larva of A. psoraliella feeds only on plants that formerly were assigned to the genus Psoralea. Specimens of A. psoraliella in the collection of the United

States National Museum of Natural History (hereafter referred to as USNM) were reared in Iowa from silverleaf Indian breadroot, Pediomelum argophyllum (Pursh) J. Grimes (labeled as

Psoralea argophylla), in Nebraska from slimflower scurfpea, Psoralidium tenuiflorum (Pursh)

Rydberg (labeled as Psoralea tenuiflora), and in Arkansas from Sampson's snakeroot, Orbexilum pedunculatum (P. Miller) Rydberg (labeled as Psoralea psoralioides (Walt.) Cory). Emergence dates of these moths are concentrated in the period extending from mid-June through early July.

Psoralidium tenuiflorum is common at a number of the Illinois prairies that I surveyed for microlepidoptera (these sites lying along a 240-km, northeast-to-southwest line extending from

Mason through Jersey counties), but no larval evidence of A. psoraliella was seen on P. tenuiflorum at these sites. This could indicate that A. psoraliella is absent from Illinois, but a more extensive survey would be necessary to assess the true status of A. psoraliella within the state.

Anacampsis wikeri Harrison, new species

(Figs. 8, 10, 12)

81

Adult (Fig. 8). Head: Smoothly scaled, uniformly brown; antenna brown, basal one-third of

each flagellomere slightly darker than apical two-thirds, giving the flagellum a faintly annulated

appearance; labial palpus brown laterally, paler and more shining brown medially; maxillary

palpus greatly reduced; haustellum clothed in pale-brown scales. Thorax: Collar; tegulae, and

dorsum of thorax uniformly brown; forewing length, 7.5mm; dorsum of forewing brown, ground

color somewhat paler than dorsum of head and thorax; sometimes with a dark-brown patch in the fold at 0.2 length of wing from base; a dusting of dark-brown scales above the fold from base to

0.6 wing length, at which point occurs a very diffuse dark-brown fascia across full width of wing, extending to 0.7 wing length, beyond which occurs an incomplete ochreous band in the form of an anterior and posterior patch, the former being the more prominent of the two; a series of five or six small dark-brown spots on margin, beginning on costa immediately beyond anterior ochreous patch and extending around apex to occupy anterior half of apical margin of wing; fringe slightly paler brown than wing, with a narrow, indistinct darker band near base of fringe; dorsum of hindwing uniformly brown, of the ground color of the forewing; fringe as in forewing; ventral surface of forewing and hindwing shining brown, slightly paler than dorsum, unmarked; ventral surface of thorax shining pale brown; all legs with coxa, trochanter, and femur concolorous with thorax, tibia and tarsus concolorous with thorax ventrally, darker and duller brown dorsally, tarsus without annulation, hind tibia with prominent brushlike fringe of narrow ochreous scales. Abdomen: Brown, darker dorsally than ventrally, both surfaces concolorous with thorax; male genitalia (Fig. 10) with valve straight, subcylindrical, length approximately

7.0x width, exceeding the combined vinculum/juxta by approximately 1.1x its length; vinculum narrowing anterad to form a narrowly-rounded V shape in ventral aspect; juxta in ventral aspect

82 with posterior margin excavated into a U-shaped curve, the posterolateral margin on either side

produced posterad into a sharp point; socius lobes prominent, reflected ventrad at a 45-degree

angle, with no excavation between the lobes in ventral aspect; gnathos an unsclerotized band

with no ventrally-projecting medial process; aedeagus with phallobase approximately 2.3x as

long as wide, occupying 0.45 aedeagus length, distal part of aedeagus slightly narrower than phallobase, with narrow lateral ridge, shallowly and evenly curved ventrad then dorsad, truncate

at apex; female genitalia (Fig. 12) with conical sclerotized structure projecting anterad from

ostium bursa, ductus bursa and corpus bursa unsclerotized, signum absent; abdominal segment 8

with middorsal posteriorly-projecting sclerotized process in lateral aspect elongate, narrow in

basal 0.45 of its length, then enlarging abruptly into a broad knob that narrows ventrally in its

apical half to form a slightly curved, thornlike apex.

Mature larva. Length, 11.5 mm; color grayish, owing to finely spiculate texture of cuticle;

head, legs, and prothoracic and anal shields blackish; sclerotized pinacula present at bases of

most of the primary setae, especially SD, L, and SV; pinacula larger on T1 and T2 than on other

segments; seta SD1 on A9 hairlike.

Life history. The species is univoltine. At the type locality in central Illinois, and

oviposition occur probably in early May. The larva feeds on leaves of leadplant, Amorpha

canescens Pursh (Fabaceae). It lives and feeds inside a shelter that it constructs by silking

together two or more leaves, usually near the apex of the plant. The larva matures during the

third week of May, after which it pupates and, in early June, emerges as an adult, in which stage

it overwinters.

83

Host plant and habitat specificity. Three species of Amorpha occur in Illinois. One of them,

Amorpha nitens Boynton, is known only from a few isolated localities (none of them prairies) in far southern Illinois (Mohlenbrock and Voigt 1959; Taft 2005) and is listed as an endangered species in the state (Herkert and Ebinger 2002). Not surprisingly, the Lepidoptera fauna (if any) that is associated with this plant in Illinois is not known at present. The remaining two Amorpha species in Illinois are widespread throughout the state (Mohlenbrock and Ladd 1978), but they occupy different habitats: A. canescens is a prairie-restricted plant, whereas Amorpha fruticosa

Linnaeus prefers wet areas (Jones 1945). A number of specialist Lepidoptera species, e.g.,

Gracillariidae: Phyllonorycter uhlerella (Fitch), feed as larvae on both of these Amorpha species, but extensive searching of A. fruticosa in Illinois by J. R. Wiker has not yielded any Anacampsis larvae. Therefore, it is concluded that A. wikeri feeds only on leadplant and thus is obligately restricted to prairie habitat.

Distribution. At present, A. wikeri is known only from Illinois prairies in Mason and Jersey

Counties. It is hoped that the present paper will encourage surveys for this moth in other areas.

Diagnosis. Externally, A. wikeri is very similar to A. psoraliella in size and coloration, to the degree that sight identification of adults of these two species that have not been reared probably is not reliable. However, A. wikeri is easily separated from A. psoraliella on genital morphology

(as discussed and illustrated herein) and on larval biology.

84 Types. Holotype male. (1) [White label]: ILLINOIS, Mason County/Revis Hill Prairie/T20N-

R7W, Section 36/May 18, 2005-Larva/Collected by James R. Wiker; (2) [White label]: Larva

found on leaves of/Amorpha canescens/May 18, 2005-J. Wiker/Pupated: Late May,

2005/Emerged: June 7, 2005; (3) [Red label]: HOLOTYPE/Anacampsis wikeri ♂/Harrison

(USNM). Allotype female. Same data as for holotype, except collected May 26, 2002, emerged

June 18, 2002; [Yellow label]: ALLOTYPE/Anacampsis wikeri ♀/Harrison.] (USNM).

Paratypes: 4 ♂♂, same data as for holotype except (1) collected May 18 2005, emerged June 6,

2005; (2) collected May 18, 2005, emerged June 9, 2005; (3) collected May 25, 2006, emerged

June 20, 2006; (4) collected May 17, 2004, emerged mid-June, 2004; 5 ♀♀, same data as for holotype except (1) collected May 18, 2005, emerged June 11, 2005; (2) collected May 26, 2002, emerged June 14, 2002; (3-5) collected May 26, 2006, emerged June 20, 2006 (three individuals)

(collection of James R. Wiker, Greenview, Illinois); also 1 ♂, Collected as larva on Amorpha canescens, USA: Illinois, Mason County, Revis Hill Prairie, 18-V-2006, T. Harrison, emerged 6-

VI-2006; 1 ♀, same data as for previous specimen, except emerged 7-VI-2006 (USNM). [All

paratypes with blue label]: PARATYPE/Anacampsis wikeri ( ♂ or ♀)/Harrison.

Etymology. Named in honor of Illinois lepidopterist James R. Wiker, who first reared this

species and brought it to the attention of the authors, and who provided much incidental

information that was relevant to the description of this moth.

85 Discussion

Species relationships within Anacampsis are difficult to evaluate, because no phylogenetic

analysis of the genus has been conducted. In the genital morphology of A. psoraliella and A. wikeri are seen two character states (reduction of gnathos in male, and elongation of A8 tergal process in female) that are unusual within Anacampsis, and which, along with similar adult coloration and shared specialization on prairie legumes as larval hostplants, might represent apomorphies that reflect recent common ancestry of these two species. This hypothesis, however, has yet to be tested in the context of a phylogenetic analysis of the entire genus.

Literature Cited

Barnes, W., and A. Busck. 1920. Gelechiidae. Contributions to the Natural History of the

Lepidoptera of North America IV: 224-230.

Clarke, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera.

Bulletin of the Brooklyn Entomological Society 36: 149-161.

Herkert, J. R. and J. E. Ebinger, editors. 2002. Endangered and Threatened Species of

Illinois: Status and Distribution, Volume 1 - Plants. Illinois Endangered Species

Protection Board, Springfield, Illinois.

Jones, G. N. 1945. Flora of Illinois. American Midland Naturalist Monograph 2: 1- 317.

Klots, A. B. 1970. Lepidoptera. Pp. 115-130 in Tuxen, S. L., ed. Taxonomist's

Glossary of Genitalia in Insects, 2nd Edition. Copehagen: Munksgaard, 359 pp.

Lee, S., R. L. Hodges, and R. L. Brown. 2009. Checklist of Gelechiidae (Lepidoptera)

86 in America North of Mexico. Zootaxa 2231: 1-39.

http://www.mapress.com/zootaxa/2009/f/zt02231p039.pdf

Mohlenbrock, R. H., and D. M. Ladd. 1978. Distribution of Illinois Vascular Plants.

Carbondale and Edwardsville: Southern Illinois University Press, 282 pp.

Mohlenbrock, R. H., and J. W. Voigt. 1959. A Flora of Southern Illinois. Carbondale and

Edwardsville: Southern Illinois University Press, 390 pp.

Pitkin, L. 1984. A technique for preparing complex male genitalia in microlepidoptera.

Entomologist's Gazette 37: 173-179.

Robinson, G. S., P. R. Ackery, I. J. Kitching, G. W. Beccaloni & L. M. Hernández, 2009.

HOSTS - A Database of the World's Lepidopteran Hostplants. Natural History

Museum, London. http://www.nhm.ac.uk/hosts. (Accessed: 20 Dec. 2009).

Taft, J. B. 2005. Conservation Assessment for Amorpha nitens Boynton. Eastern

Region of the Forest Service, Threatened and Endangered Species Program,

Technical Report 2005 (6): 1-35.

USDA, NRCS. 2009. The PLANTS Database (http://plants.usda.gov, 5 November 2009).

National Plant Data Center, Baton Rouge, LA 70874-4490 USA.

Figures

Figures 7, 8. Adult moths. 1, Anacampsis psoraliella; 2, Anacampsis wikeri.

Figures 9, 10. Male genitalia. 3, Anacampsis psoraliella; 4, Anacampsis wikeri; a, tegumen,

gnathos, and uncus, ventral aspect; b, valvae, juxta, and vinculum, ventral aspect; c, valvae and

87 vinculum, right-lateral aspect; d, aedeagus, left-lateral aspect. Differences are seen in respective forms of the valve, juxta, vinculum, and aedeagus.

Figures 11, 12. Female genitalia (abdominal segments 7 and 8, right-lateral aspect). 5,

Anacampsis psoraliella; conical invagination absent at ostium bursae, A8 tergal process elongate, straight, and dorsoventrally flattened along entire length; 6, Anacampsis wikeri; conical invagination present at ostium bursae, A8 tergal process elongate, subcylindrical, gently curved ventrad from base to 0.6 its length then abruptly doubly recurved and enlarged then narrowed into knoblike formation with thornlike apex.

88

89

90 Author’s Biography

B. S., Zoology, Eastern Illinois University

M. S., Entomology, University of Illinois at Urbana-Champaign

Formerly Research Specialist, Department of Entomology, University of Illinois at Urbana-

Champaign, where he was involved in molecular characterization of coding for cytochrome oxidase enzymes in larval black , polyxenes. During his time as Research Specialist, he developed an interest in biotaxonomy of North American microlepidoptera. He has authored and coauthored a number of papers on microlepidoptera, these including the descriptions of several previously unknown or unrecognized Nearctic species, and he is in the process of developing a web site on microlepidoptera of eastern North America, with emphasis on Illinois (http://www.microleps.org). For his doctoral dissertation (Department of Entomology, University of Illinois at Urbana-Champaign), he implemented his interest in moths in a study in which he compared microlepidoptera of Illinois hill prairies, evaluated microlepidoptera families for likelihood of their harboring prairie-restricted species, analyzed moth diversity in biofuel crops versus native prairie, and described two new species of prairie- restricted microlepidoptera from Illinois.

91