The Immature Stages of the Rose Curculio Meryhynchites Bicolor (Fabricius) (Coleoptera: Rhynchitidae): with Notes on Its Biology in Northern Illinois

Loyola University Chicago Loyola eCommons

Master's Theses Theses and Dissertations

1979

The Immature Stages of the Rose Curculio meryhynchites Bicolor (Fabricius) (Coleoptera: Rhynchitidae): With Notes on Its Biology in Northern Illinois

Sheila S. Kuritsky Loyola University Chicago

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Recommended Citation Kuritsky, Sheila S., "The Immature Stages of the Rose Curculio meryhynchites Bicolor (Fabricius) (Coleoptera: Rhynchitidae): With Notes on Its Biology in Northern Illinois" (1979). Master's Theses. 3095. https://ecommons.luc.edu/luc_theses/3095

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This work is licensed under a Creative Commons Attribution-Noncommercial-No Derivative Works 3.0 License. Copyright © 1979 Sheila S. Kuritsky THE IMMATURE STAGES OF THE ROSE CURCULIO, MERHYNCHITES BICOLOR (FABRICIUS) (COLEOPTERA:RHYNCHITIDAE), WITH NOTES ON ITS BIOLOGY IN NORTHERN ILLINOIS

by Sheila s. Kuritsky

A Thesis Submitted to the Faculty of the Graduate School of Loyola University of Chicago in Partial Fulfillment of the Requirements for the Degree of Master of Science December 1979 ACKNOWLEDGMENTS

I would like to thank the following people for their assistance in the preparation of this thesis, for without them this project could not have been completed: my committee members, Drs. Jaskoski and Peluso, Loyola University of Chicago, for their valuable suggestions; and Terry N. Seeno, California Department of Food and Agriculture, for loan of specimens. Special thanks to Dr. Rouffa, University of Illinois at Chicago Circle, for allowing me access to the James Woodworth Prairie Preserve, where the life cycle of Merhynchites bicolor was studied and immature stages obtained. I am most grateful however, to my adviser, Dr. Robert W. Hamilton, Loyola University of Chicago, not only for his infinite patience and assistance, but also for enlightening me about the wonderful world of weevils.

ii VITA

The author, Sheila Sue Kuritsky, is the daughter of Morris Murray Kuritsky and Flora (Dubin) Kuritsky. She was born June 1, 1953, in New York, New York.

Her elementary education was obtained in the public schools of Oradell, New Jersey, and secondary education at River Dell Regional High School, Oradell, New Jersey, where she graduated in 1971.

In September, 1971, she entered Boston University, and in May, 1975, received the degree of Bachelor of Arts with a major in biology. Her pertinent undergraduate coursework was Botany, Inorganic Chemistry, Organic Chemistry, Histology, Calculus, Microbiology, Parasit ology, Genetics, Human Ecology, Physics and Spanish.

In September, 1977, she entered the Biology Graduate Program at Loyola University of Chicago. Her pertinent graduate coursework was Entomology, Limnology, Evolution, Medical Entomology, General Physiology and Developmental Biology. In January, 1978, she was granted a teaching assistantship. iii In 1978, she also became a member of the Illinois Academy of Science and the Entomological Society of America. During her graduate studies she attended two North Central Branch meetings of the ESA, two national meet~ngs of the ESA, and presented a paper entitled "The Biology of Merhynchites bicolor (Fabr.) in Northern Illinois" to the ESA.

iv TABLE OF CONTENTS Page

ACKNOWLEDGMENTS • • • • • • • • • • • • • • • ii

VITA • • • • • • • • • • • • • • • • • • • • iii

LIST OF FIGURES . . • • • ...... vi LIST OF ABBREVIATIONS • • • • • • • • • • • • viii

LIST OF TABLES • • • • • • • • • • • • • • • • xi

INTRODUCTION • • • • • • • • • • • • • • • • • 1

REVIEW OF RELATED LITERATURE • • • • • • • • • 3 Taxonomy • • • . • • • • • ...... 3 Economic Importance • • • ...... 4 Control ...... • ...... 6 Larval and Pupal Terminology ...... ? MATERIALS AND METHODS ...... 9

RESULTS AND DISCUSSION • • • • • • • • • • • • 13 Life Cycle Observations . • • • • • • • • • 13 Larval Description . . . • • • • • • • • • 1? Pupal Description • • • • • • • • • • • • • 21 Diagnosis • • . . • • • • • • • • • • • • • 23

FIGURES • • • • • • • • • • • • • • • • • • • 2?

TABLES . . • • • • • • • • • • • • • • • • .53 LITERATURE CITED ...... 5.5

v LIST OF FIGURES Figure Page 1. Adult of M. bicolor on bud of Rosa carolina L. • • • • • . . ~ . . . 27 2. Damage to rose bloom by adult of M· bicolor • • • • • • • • • • • • 27 3· Oviposition-punctured rose hip • • • • • 27

4. Egg of M. bicolor within rose hip • • • • 2?

s. Larva of M. bicolor within rose hip • • • 27 6. Larval feeding damage to rose achenes • • 27

7· Plastic crisper • • • • • • • • • • • • 27

8. Plaster rearing cast • • • • • • • • • . 27

9. James Woodworth Prairie Preserve . • • • 29

10. Rose bush plastic wrap • • • • • • • • 29

11. Excavation of test plot • • 0 • • • 0 0 29

12o Lining of test plot 0 0 • 0 0 0 • . • • 29

13. Finished test plot • • • . • • • • • • 29

14. Larva of M· bicolor • • 0 • • • • • • • 31

15. Head of Mo bicolor larva • . • • • • • • 31

16. Head of M· bicolor larva • • • • . • • • 33

17. Head of M. bicolor larva . • • • • • • • 33.

18. Left antenna of Mo bicolor larva • • • • 35

19. Labrum of M. bicolor larva . . • • • • 35

20. Labrum-epipharynx of M· bicolor larva • 0 35

21. Left mandible of M· bicolor larva • • • • 37

22. Labium of M. bicolor larva . . • 0 • • • 37 vi Figure P.age

23. Left maxilla of M· bicolor larva • • • • 39 24. Prothoracic spiracle of Mo bicolor larva o • o • • • • • • • 0 0 39 25. Thorax and abdominal segments I and II of Mo bicolor larva 0 . . • 0 . • • • 0 0 41 26o Thorax and abdominal segments I and II of M· bicolor larva 0 . 0 0 . 0 • . • 41 27. Abdominal segments VII-X of M. bicolor larva • • . • • • • • 0 • • • 43

28. Pupa of M. bicolor . • 0 0 0 0 • • 0 0 0 45

29o Pupa of Mo bicolor 0 . • • • 0 0 . • 45 JO. The effect of temperature on larval emergence, 1978 • 0 • • . 0 • • . 47 31. The effect of temperature on larval emergence, 1979 • . . • • • . • • 49 )2. The effect of temperature on pupation, 1979 • • 0 • • 0 • • • • . • 51

vii LIST OF ABBREVIATIONS - LARVA OF M. BICOLOR

Abl to AbX, abdominal segments acap, accessory sensory appendage of antenna AlA, alar area als, anterolateral setae of e~ipharynx ams, anteromedian setae of ep~pharynx AnL, lobes of anus Ant, antenna Anu, annulus of spiracular air tube art, basal article of antenna ATb, air tube of spiracle asp, asperities cd, cardo of maxilla Cl, clypeus des, dorsal epicranial setae D~t, pronotum or dorsum of prothorax Enc, endocarina EPl, epipleurum ES, epicranial suture EuSt, eusternum Fldl to FldiV, folds of abdominal segments Fr, frons fs, frontal setae Lb, labium LbP, labial palpus les, lateral epicranial setae Lm, labrum LmR, labral rod lms, labral setae lsl, lateral sensillum of labrum Ma, mala gf maxilla Md, mandible mds, mandibular setae msn, mandibular sensillum msp, median spines of epipharynx Mx, maxilla MxP, maxillary palpus PdA, pedal area pes, posterior epicranial setae viii Pl, pleurum pla, placodeum pms, postmental setae PMt, postmentum PrD, prodorsum PrmS, premental sclerite Prt, peritreme of spiracle PsD, postdorsum Sl, sternellum sn, sensillum snp, sensory pores of epipharynx Sp, spiracle SpA, spiracular area sps, subcutaneous pigment spots St, stipes of maxilla Stn, sternum Thi to Thiii, thoracic segments

ix. LIST OF ABBREVIATIONS - PUPA OF M. BICOLOR

Head and Rostrum - baR, basirostral setae diR, distirostral setae subO, suborbital setae Prothorax - alP, anterolateral pronotal setae amP, anteromedian pronotal setae mP, median pronotal setae mlP, mediolateral pronotal setae plP, posterolateral pronotal setae pmP, posteromedian pronotal setae Mesonotum - msN, mesonotal setae Metanotum - mtN, metanotal setae Abdomen - dsT, discotergal setae ltS, laterosternal setae ltT, laterotergal setae PP, posterior processes sltS, sub-laterosternal setae

X LIST OF TABLES Table Page 1. Summary of Selected Larval Characters . . • • . • • • • • '" . • • 53 2. Summary of Selected Pupal Characters . . . • • • • . . . . • • • 54

xi INTRODUCTION

Merhynchites bicolor (Fabricius) is a member of the family Rhynchitidae, the tooth-nosed snout beetles. There are 44 recognized species in this family in America north of Mexico. The adults of this group are morphologically distinct from all other weevils in that they have straight antennae with a )-segmented club, their mandibles are toothed on the outer and inner surfaces, they have dis tinct parts to their maxillae, a 4-segmented maxillary palpus, )-segmented labial palpi, and no distinct labrum {Hamilton, 1969). However, many authors do not recognize the family Rhynchitidae (Van Emden, 19)8, Peterson, 1960, Kissinger, 1964, et. al.), but place this species and others in the subfamily Rhynchitinae. Kissinger was the last author to treat the family Curculionidae as a whole, with Rhynchitinae as one of 42 subfamilies. No detailed work has ever been done on the immature stages of this weevil to support their placement in this family. The principal goal of this project was to describe and illus trate the immature stages of the rose curculio to show that it is a rhynchitid with unique rhynchitid characteristics, and therefore morphologically distinct from all other groups of weevils. At the present time, a Coleoptera catalogue is being prepared by the USDA in which 12 fam-

1 2 ilies of weevils will be recognized for America north of Mexico. More data is needed to support the elevation of this group of weevils to family instead of a subfamily of Curculionidae and to verify its status in the USDA cata logue. M. bicolor is a holometabolous and univoltine insect in northern Illinois. It is ope of two rhynchitids whose economic importance is known in the literature. The adult rose curculio is a pest of roses, feeding on the develop ing buds and ovipositing into the hypanthium. Eggs hatch and the larvae feed on and develop within the hypanthium until the fall. At this time, mature larvae leave the hips, drop to the ground, burrow into the soil and form a cell in which they diapause. Pupation occurs in late spring, with adult emergence shortly thereafter. While rearing the rose curculio to obtain the immature stages, available life cycle data was verified. REVIEW OF RELATED LITERATURE

TAXONOMY

Merhynchites bicolor (Fabricius) was first des cribed as Curculio bicolor Fabricius in 1775. In 1792, Fabricius also listed it in the genus Attelabus Linnaeus. Olivier (1807) realized that it did not belong in either genus and transferred it to Rhynchites Schneider, 1791. The genus Merhynchites was first erected by Sharp (1889) and included only two species--Rhynchites bicolor (F.) and Rhynchites hungarius Herbst, the latter does not occur in North America and is now placed in Involvulus Schrank. In 1912, Cockerell recognized a race that differed from R. bicolor (F.) in that it had a black head, less coarsely sculptured elytra, and the elytra were lacking rows of evident coarse punctures. He named this form Rhynchites bicolor wickhami Cockerell. In 1913, Pierce designated R. bicolor (F.) as the genotype for Merhynchites and listed six color varieties of the species, which are endemic to North America. Kissinger (1964) listed R· bicolor as one of 29 species in the genus Rhynchites. Hamilton (1969) recognized four of Pierce's six varieties as valid species, M. bicolor (F.), Merhynchites wickhami (Cockerell), Merhynchites tricarinatus (Green), and Merhynchites palmi (Schaeffer). Characteristics which delimit the genus 4 Merhynchites are given by Hamilton (1971).

ECONOMIC IMPORTANCE

The economic importance of M. bicolor as a pest of roses was first recorded by Harris (1862). He observed damage done to both cultivated and wild rose bushes by adults and larvae of the genus Rhynchites. Harris also gave a brief description of the adult weevil. Further comments were made on the "rose curculio" and its economic importance in damaging roses, blackberries, and raspberries (Chittenden, 1901). Fall (1901) stated that R. bicolor was found wherever there were wild roses. Gates (1909) was the first to observe and describe damage done to wild and cultivated roses in Massachusetts. He observed that flower buds and leaves of the Japanese rose, Rosa rugosa Thunb., were damaged byE· bicolor. Dickerson (1910) elaborated on the damage done to R. rugosa in New Jersey by this weevil. In addition, Dickerson added information on its behavior in that females oviposit into the "seed capsule". He also observed larval emergence from the hips in the fall, but made no mention of where overwintering occurred. The rose curculio was listed among seven species of Rhynchites and a brief description of the adult and its North American distribution was provided (Blatchley, 1916). Blatchley also commented that the larvae develop in the "hips", and that the adult not only punctured E· rugosa 5 buds, but also had caused blackberry fruits to decay due to the feeding punctures. Essig (1958) elaborated on the description of the eastern form of the adult, and also listed various western color phases and their ranges. He also noted that all species fed on wild and cultivated roses, although they have been observed on other host plants such as raspberry, blackberry, and thimbleberry. Essig observed that oviposition normally occurred in the rose hip or ovary and that overwintering took place in the soil. R. bicolor was listed as an inquiline with the thim bleberry gallmaker, Diastrophus kincaidii Gillette (Hymenoptera:Cynipidae) (Wangberg, 1975). The western form of the rose curculio was first men tioned by Cassidy (1888) as a pest of roses and raspberries in Colorado. The "rosebud curculio" was observed on wild and cultivated roses in Montana (Cooley, 190J). Eggs were not only found in rose hips, but also in buds. Cooley also stated that the weevils bored into the stem of the rose causing the bud to wilt and probably drop off. Lovett (1915) verified that weevil damage occurred in Oregon. He also confirmed that buds contained eggs and the stem was punctured, causing the bud to droop and die. Lovett also stated that larvae were never found in the buds, but instead in the rose seed pods. Gillette and List (1921) were the first to call the western rose snout beetle Rhynchites wickhami Cockerell and stated that it can be 6 distinguished from the eastern form by its black head. In Manitoba, the rose curculio laid eggs in the buds and the larvae then moved to the hips to feed (Robertson, 1924). Robertson also stated that mature larvae (not pupae, as was previously believed by Cooley, Lovett, et. al.) over wintered in the soil. Hoerner (1936) published the dis tribution for the western rose curculio, Rhynchites bicolor wickhami Ckl. He also verified that eggs were laid in the rose buds and that the stem is then punctured, providing a dead bud for larval development. Hoerner pro posed that pollen is probably the food for the early larvae. W.V. Balduf (1959) observed that Rhynchites bicolor was one of four insects that actually fed on rose hips. He observed that the eggs were oviposited into the hips and after hatching, the young larvae fed on the achenes until full grown. The mature larvae then emerged from the hips, dropped to the ground, and burrowed from 1-4" to overwinter. Balduf concentrated his work on identifying all the insects that inhabit rose hips, and he stated the need for greater concentration on each individual insect.

CONTROL

Cooley (1903) was the first to suggest the control method of handpicking or spraying adult rose curcul.ios with Paris green, and as a last resort to protect cultiv- 7 ated roses, he suggested destroying wild roses where the beetles bred. Lovett (1915) suggested that adults should be jarred from the plants and destroyed. He also stated that this control should be supplemented with eradication of wild roses, handpicking injured "seed pods", spraying lead arsenate when the adults emerged, and spraying white hellebore when the blossoms emerged. Hellebore is an alkaloid derivative of the roots of white "false hellebore" plants, Veratrum album, from Europe. Blatchley (1916) also recommended handpicking or spraying with Paris green for adult control. Robertson (1924) also mentioned hand picking adults, spraying with lead arsenate, and destroy ing all hips late in the summer, before the larvae had left them. Handpicking adults and injured hips, dusting with calcium arsenate for adults, and applying carbon disulfide emulsion for overwintering larvae in the soil were suggested as additional control methods (Hoerner, 1936). Essig (1958) obtained good control over the weevil with a 5% nicodust insecticide.

LARVAL AND PUPAL TERMINOLOGY

Van Emden (1938) dealt with the taxonomy of eight subfamilies of Rhynchophorous larvae and published a key to the main groups and species. The genus Pissodes Germar was used to demonstrate basic terminology for weevil larvae (W.H. Anderson, 194?). Crowson (1955) listed characteristics 8 present in larvae of the family Curculionidae and its sub families in his natural classification of families of Coleoptera. D.M. Anderson (1968) found differences in the pupae of two subspecies of Anthonomus grandis and also found anatomical differences in the sexes of these pupae, which he proposed were consistent for all weevils. A terminology using chaetotaxy and other pupal structures was developed and applied to 47 species of anthonomine weevil pupae (Burke, 1968). MATERIALS AND METHODS

This project began by consulting the available life cycle data of Merhynchites bicolor. The collection data on specimens in the Loyola University of Chicago collec tion and the private collection of Dr. R.W. Hamilton revealed that adults could be collected on wild roses early in June (Fig. 1). At this time, extensive feeding damage to rose blooms was observed (Fig. 2). On June 28, 1978, adults of M. bicolor were collected from the James Woodworth Prairie Preserve in Niles, Illinois (Fig. 9) and a relatively undisturbed prairie area along the Soo Line railroad tracks in Mount Prospect, Illinois. These adults were handpicked from wild rose bushes and placed in plastic crispers with a thin layer of bottom sand

(Fig. 7). Fresh bud~ and hips of the host plant, Rosa carolina L. were placed in the sand daily, providing the adults food and oviposition sites. Copulation was also observed in the crispers. Hips with oviposition punctures were removed and the eggs and larvae within were preserved. KAAD (kerosene, alcohol, acetic acid and dioxane) or FAA . (formaldehyde, acetic acid and alcohol) were used as fixa tives, and after 24 hours, the eggs and larvae were then transferred to 70% ethyl alcohol.

9 10 At the two collection sites, rose hips with ovi position punctures were collected from June through Nov ember, 1978, and at two-week intervals from July through September, 1979 to determine that the behavior observed in the laboratory also occurred naturally in the field (Fig. J). Some eggs and larvae were periodically dissected out of these field-collected hips and compared with the laboratory-reared eggs and larvae (Figs. 4 & 5). The majority of these hips, however, were not dissected but instead, were kept on sand in plastic crispers so that the larvae inside could mature. On October 7, 1978 and September 1, 1979 Woodworth Prairie rose bushes were also wrapped with plastic sheet ing and tied below so that larvae emerging from the hips would be trapped (Fig. 10). Bags were checked regularly and emergence dates recorded. Some larvae trapped by the plastic wrapping were removed and allowed to burrow into potting soil in large, wide-mouthed jars. These were kept outside for the winter and excavated in the spring to deter mine depth of larval burrowing. Other larvae were kept in refrigerators at approximately 4°C in plaster rearing casts filled with potting soil (Fig. 8). These plaster casts were mainly used to prevent larval dessication dur ing diapause. The centerwell portion of the cast was covered with glass and a 5 em X 2 em well was watered regularly. In this way, the soil was kept relatively moist without disturbing the larvae. Refrigerated speci- 11 mens were removed and placed outside in early spring in order to induce pupation. Pupae were fixed using either KAAD, FAA or by dropping them into boiling water. Those fixed using the latter method were transferred to 70% ethyl alcohol after 5 minutes, while the others were transferred after 24 hours. Larvae were also metered under natural soil condi tions at the Woodworth Prairie site. In October 1978, a J-foot square area was excavated (Fig. 11). Plastic screening was laid 1 foot underneath the ground surface, effectively lining the excavation {Fig. 12). The plot was then filled in with prairie soil. Approximately 200 hips that were believed to be oviposition-punctured were collected and placed on the surface on the plot, in anti cipation of concentrating emerging larvae in this plot. The hips were then covered with a heavy hardware cloth to prevent consumption by other animals (Fig. 1J). This plot was excavated in May 1979 and data on the larvae and pupae in their natural habitats was compared with the data obtained in the laboratory. All illustration and measurements were made using an ocular reticule. The finished drawings were made in India ink and reproduced as plates. The abbreviations used are in accordance with Snodgrass (1935), Anderson (1947), Burke (1968) and Anderson (1968). Larval head 12 capsules and mouth parts were dehydrated in ethyl alcohol, transferred to xylene and mounted in piccolyte media on slides. Head capsules were cleared in KOH (potassium hydroxide) prior to mounting to remove musculature •

• RESULTS AND DISCUSSION

LIFE CYCLE OBSERVATIONS

Behavior of M. bicolor was observed- at both Illinois field sites and also in the laboratory. Copulation was observed on the host plant, Rosa carolina. Oviposition was best observed in the laboratory, since this behavior may take up to 15 minutes. Oviposition that was observed agreed with that described by Dickerson (1910). Oviposi tion was initiated by the female excavating an egg pit in the hypanthium with her beak, with the excavation usually ending up as deep as the length from her beak to the eyes. The female then turned around and deposited an egg in the pit. She then completed the behavior by turning around again, pushing the egg down into the pit with her beak, and covering the opening with what is probably a salivary secretion (Balduf, 1959). The presence of this black covering almost always indicated the presence of an egg or larva within the hip. Most of the eggs that were recov ered from the hips were found deep in the achene cavity. This oviposition behavior puts the young larva in close proximity to its food supply. The egg of M. bicolor was slightly ovoid and dirty white when first oviposited. They measure 0.84 : 0.07 mm in length and o.6J : 0.10 mm in width (N = 5 eggs) and as lJ 14 they mature, they yellow and the mandibles of the larvae become visible through the chorion. Eggs oviposited in the laboratory hatched within 14 days. After hatching, the young larvae began feeding on the dicotyledons within the achenes (Fig. 6). The early instars were usually found within a single achene, hidden beneath large amounts of brown frass. Achenes were entirely bored out by the larvae, leaving the shells intact. The maturing larvae soon become so large that they no longer occupy single achenes. Instead, they were usually found within the achene cavity in which the frass continued to accumulate. With the onset of cooler weather, the mature larvae left the achene cavity and bored through the hypanthium, emerging to overwinter in the soil beneath the rose bushes. Larvae probably burrow into the soil immediately after emerging, but this was not observed in the field. However, emerging larvae that were caught in the plastic wrapping at the Woodworth Prairie burrowed immediately when placed on soil in the plaster rearing casts. Temperature appeared to be the primary stimulus for larval emergence from the hips (Figs. JO and 31). The majority of larvae were observed to emerge from the hips when temperatures began to drop in the fall. This obser vation agreed with those of Balduf, who found that 88% of his larvae emerged when daily low temperatures were in the J4-54°F range. 15 Depth of larval penetration varied with the con sistency of the soil (Balduf, 1959). Larvae allowed to burrow into potting soil penetrated easier and were thus located deeper (usually 4") than those that were allowed to burrow into prairie soil. Some larvae that were allowed to burrow were later excavated and found inside smooth oval cells in the soil. It is in these cells formed by the larvae that they diapause until pupation the following May.

In ~orne cases, parasitism of M· bicolor larvae by Luchatema baldufi Walkley (Hymenopteratichneumonidae) caused diapause to extend beyond the normal pupation time (Balduf, 1959). This phenomenon of prolonged diapause was observed at the Woodworth Prairie. On June 1, 1979, two mature larvae of a parasite emerged from two weevil larvae that had failed to pupate. These two ichneum6nids came to the surface of the soil and began to spin cocoons; however, they did not successfully complete their cocoons and soon died. As a result, these two parasites were never positively identified as 1· baldufi. However, it is highly probable that they indeed were this ichneumonid because at this time in June, adults appeared in good numbers at the Woodworth Prairie on rose bushes. During the summer of 1979, M. bicolor larvae were observed to be heavily parasitized by a hymenopteran that was not recorded by Balduf. This unidentified chalcid 16 parasite is a solitary ectoparasite. In many instances, dead and shrunken weevil larvae were found inside an oviposition-punctured hip. In two instances, pupae of the parasite were found in the achene cavity alongside the weevil remains and frass. Pupation was successfully induced in the laboratory by removing five larvae from the refrigerator in late March and early April. After J-4 weeks of exposure to room temperature, these five larvae pupated. In the field, however, M· bicolor larvae free from parasites pupate in mid-May. The end of diapause is marked by the exsertion of the larval head. On May 18, 1979, the test plot at the Woodworth Prairie was excavated. The soil was hand sorted, and two larvae and two pupae of M· bicolor were recovered. This low recovery rate of larvae and pupae from the test plot was probably due to a high rate of parasitism by a number of species of hymenopterous parasites. However, since pupae were recovered, the pupation period had begun and probably continued until early June (Fig. J2). The period of adult emergence was expected to begin early in June, and the first adult was observed on the host plant at the prairie on June 4, 1979. Balduf (1959) observed an overall adult season of about 80 days at Urbana, Illinois. This period was cal culated from the onset of pupation until the last date 17 that an adult was observed. At the Woodworth Prairie, which is considerably north of Urbana, a shorter adult season would be expected. Observations over two summers gave an overall adult season of 68 days. -

LARVAL DESCRIPTION

Body: (Fig. 14) Cyphosomatic, off white, with numerous transverse folds. Length 4.9-6.6 mm. Asperities confined to dorsum of body, generally distributed where setae are located. Head: (Figs. 15, 16, 17) Retracted, brown anteriorly, off white posteriorly, wider than long. Head capsule width 0.85-0.90 mm. At anterior end of frons, there is a sinuous band of pigmentation. Ocelli absent. Three pairs of obvious subcutaneous pigment spots present; the first and largest pair is located posterolateral to the antennae. The second and third pairs of pigment spots are smaller and more subcutaneous than the first pair. There are 2 more pairs of less conspicuous pigment spots that are located near the lateral epicranial setae. Antenna (Fig. 18) consisting of one membranous article bearing a conical accessory appendage and five small setae. Catapophyses distinct. Hypopharyngeal bracon present. Frontal suture indistinguishable. Epicranial suture less than ~ as long as head and incomplete anter- •

iorly. Endocarina ~ as long as frons. Frons with 4 pairs of setae, fs -fs long and located at the posterior 3 5 margin of the frontal pigmentation band. Fs absent. Fs 1 2 is short and located posterior to fs . Three pairs of 3 dorsal epicranial setae, des1 and des2 at frontal suture and des near posterior pigmen~ spot. Three pairs of 3 lateral epicranial setae, the anterior pairs long and the posterior pair short. Four pairs of minute posterior epicranial setae present. Three pairs of ventral epi cranial setae, the two anterior pairs long and the pos terior pair short. Clypeus translucent with 2 pairs of setae, short and subequal. Labrum (Fig. 19) translucent, its anterior margin broadly rounded. Four pairs of labral setae, lms4 short. There is a pair of lateral sensilla near the base of lms2 . Labral rods elongate and parallel. Epipharynx (Fig. 20) with J anterolateral setae (one side only), 2 anteromedian setae (total number), and 2 median spines (total number). There are 5 sensory pores between the anteromedian setae and the median spines and 6 sensory pores behind the median spines. Epipharynx without asper ities. Mandible (Fig. 21) with two apical teeth, a small basal tooth, one sensillum, and 2 subequal setae. Labium (Fig. 22) with 2-segmented palpi, each article bearing a placodeum. Terminal article with 5 minute papillae. Pre mental sclerite complete. Prementum with 1 pair of setae and 2 pairs of placodea. Mesal aspect of ligula with 1 19 pair of sensory structures and 2 pairs of setae. Post mentum with J pairs of setae, all located at the lateral margins of the labium, with a placodeum between pms and 1 pms2 • Maxilla (Fig. 23) with )-segmented palpus, the second and third articles each bearing a short seta. Each article bears a placodeum. Maxillary stipes with J setae, J·peg-like sensilla and 1 spine-like sensillum. Mala with 5 parallel-sided, blunt-tipped sensilla, 4 blade-like, pointed sensilla, J spine-like sensilla and 1 setose sen sillum on the mesal margin. Thorax: (Figs. 25 and 26) (Setae described on one side of body only) Pronotum with 13-16 setae, the 5 along the posterior margin borne within a prominent band of asperities. Thoracic spiracle (Fig. 24) bicameral and annulated. Spir acular area of mesothorax with 5 setae, 2 long and J short. Spiracular area of metathorax with J setae. Prodorsa of mesothorax and metathorax with J short setae. Postdorsa of mesothorax and metathorax with 4 long setae arranged in a row, with a variable number of short setae, usually 5, interspersed between the 4 long ones. Alar areas of mesothorax and metathorax with 2 setae, one short, the other as long as postdorsal setae. Propleurum with 9 setae. Epipleura of mesothorax and metathorax with 9 setae. Pleura of mesothorax and metathorax with 1 long seta. Pedal areas of all thoracic segments multisetose, with numerous long setae. Presternum multisetose, meso- 20 sternum with 12 setae, and metasternum with 6 setae. Abdomen: (Figs. 25 and 26) (Setae described on one side of body only) A pair of spiracles present on each of the first 8 segments. Spiracles all lateral, bicameral, length of air tubes equal to the diameter of the peritreme. Air tubes directed dorsad within each segment. Spiracles lacking any sclerotized areas dorsoposteriorly. Typical abdominal segment with J folds, with fold III reduced dorsally and developed laterally. Fold I is absent. Fold III with 5 short setae. Prodorsum of typical abdominal segment with 4 short setae. Postdorsum of typical abdom inal segment with 4 long setae arranged in a row, with a variable number of short setae, usually 6, interspersed between the 4 long ones. Spiracular setae absent. Epi pleurum of typical abdominal segment extends dorsad in front of spiracle. Epipleurum with 2 conspicuous setae and a variable number of short setae. Pleurum of typical abdominal segment with 4-6 setae, 2 of which are usually longer than the others. Pedal area of typical abdominal segment with 2 setae, 1 short, the other long. Eusternum of typical abdominal segment with J short setae. Ster nellum present. Abdominal segments VII and VIII (Fig. 27) similar to typical abdominal segment, except that areas are less distinct and setae are more crowded. Anus ter minal and surrounded by four lobes, the lateral lobes with 2 setae each. '21 Material examined: Ten larvae, reared in the laboratory or field collected.

PUPAL DESCRIPTION Body: (Figs. 28 and 29) Length 5.75-6.75 mm. Rostrum: Two pairs of distirostral setae, borne on tub ercles. One pair of minute setae at extreme rostral apex. One pair of basirostral setae, located half way between the developing eyes and the antenna! bases. Head: Two pairs of suborbital setae located at the base of each developing eye, the medial pair of these ! the length of the lateral pair. Supraorbital and frontal setae both lacking. Prothorax: Prothoracic depressions absent. All pronotal setae are associated with tubercles; the setae are all of approximately the same length. Two pairs of anteromedian setae are borne apically on large tubercles. Five pairs of anterolateral setae are present. There are usually five pairs of mediolateral setae, but in some pupae, some of these pairs were lacking. These mediolaterals are grouped between the anterolaterals and the posterolaterals. One pair of median setae borne on widely separated tuber cles. At the posterior margin of the pronotum are five pairs of posteromedian setae, arranged in a row, some of the tubercles being fused. Two pairs of posterolateral setae are located singly toward the lateral edge of the '22 pronotum. Mesonotum: Four pairs of mesonotal setae all borne on tubercles which are clustered in a curved row. Antero notal setae are lacking. Metanotum: Four pairs of metanotal setae like the meso notals, except that the setae in each curved row are more widely separated. Anteronotal setae are again lacking. Legs: Five non-tuberculate setae at distal ends of fem ora of each leg. There is also a prominent tuberculate process on each terminal metathoracic tarsomere. Abdomen: Five pairs of discotergal setae on each of the first eight terga. All setae of approximately the same length and borne on tubercles. There are numerous very small setae anterior to and interspersed between the dis cotergals on each abdominal segment, increasing in number posteriorly. There are also a variable number of minute setae in the anterior tergal area of the posterior abdom inal segments. Anterotergal setae are lacking on all seg ments. Five pairs of laterotergal setae on segments 4-8, 2 pairs conspicuous and subequal, the other 3 pairs small. Laterotergals are minute or lacking on abdominal segments 1-J. Spiracles only on the first 6 segments. Two pairs small laterosternal setae are usually present, but on some pupae, there were more. Two pairs of minute sub-latero sternal setae. Segment 9 lacks setae, but does bear a pair of long, pointed, sclerotized posterior processes 23 that are widely separated and curve dorsally. Material examined: Six pupae, reared in the laboratory or field collected.

DIAGNOSIS Van Emden (1938) lists characteristics found in the larvae of Rhynchitinae, Rhynchitini, Rhynchites, and Rhynchites (Merrhynchites) bicolor F. Most of these characteristics are present in the larvae examined in this study. Van Emden states that all rhynchitines have long epipharyngeal rods, a retracted head, distinct endo carina, no palpiger (a labial lobe which bears the pal pus), thoracic spiracle on prothorax, abdominal tergites with 2 transverse folds, and two anteromedian epipharyn geal setae. I found all of these characters to be pre sent in the larvae of M. bicolor. Crowson (1955) includes the rhynchitids in the fam ily Attelabidae. He characterizes the larvae of the fam ily as having the prothorax overlapping the vertex of the head, 2-segmented labial palpi, 2 basal sensillae on the labrum, frontal sutures reaching the mandibular articu lations, and abdominal segmer1ts with 2 dorsal folds. Most of these characters are present in the M. bicolor larvae with the exception of the basal sensillae and the complete frontal sutures. Crowson states that the rhynchitine larva has a separate prementum and mentum and a thoracic spiracle 24 in the prothorax. These characters are present in the M. bicolor larvae, but they are also present in other cur culionid subfamilies, therefore they are not good diagnos tic characters at the family or subfamily,level. Peterson (1960) lists the following characteris tics in his description of larvae of the family Curcul ionidae: larvae usually cyphosomatic, body segments with J or more plicae dorsally; plant feeders usually light colored with a pigmented head, which is retracted in the Rhynchitinae; hypognathous mouthparts, distinct frons, more than 2 pairs of pigment spots in Rhynchitinae, very small antennae, mandible without molar area, maxilla usually with 2-segmented palpus, labial palpi with 2 seg ments, )-segmented thorax without legs, and conspicuous spiracles. Some of these characters are present in the larva of M. bicolor, although there are only 2 trans verse abdominal folds, an indistinct frons, a conspicuous antenna and a )-segmented maxillary palpus. Therefore, it appears that the following characters are apparently unique to the rhynchitids: two or more sub cutaneous pigment spots, retracted head, !-segmented con spicuous antennae, indistinct frons, long labral rods, )-segmented maxillary palpus, 2 anteromedian epipharyngeal setae, and 2 transverse abdominal folds (Table 1). Characteristics assigned to the tribe Rhynchitini by Van Emden are: three-segmented maxillary palpus, first 25 abdominal fold of segments 1-4 not more convex than pos terior fold, distinct asperities, 2 exterior epipharyn geal anterolateral setae at exterior part of front mar gin and side by side, ninth abdominal tergites and ster nites of equal length, anus terminal and X-shaped. All of these characters were found to be present in the larvae of M· bicolor. Characteristics assigned to the genus Rhynchites by Van Emden are: anterior transverse folds of abdomen without ampullae (swellings), moderately defined ventral lip of anus, mandible with small sub-basal tooth, labial palpi separated by 2 times their width, sclerotization of mentum narrower than that of prementum, setae not arising near it, and dorso-interior row of maxillary spines regular. All of these characters were found on the larvae that were observed, with the exception of the mentum sclerotization. None of the larvae that were observed had sclerotization on the mentum of the labium. Lastly, Van Emden states that these characters are found in the species~· (Merrhynchites) bicolor F.: spiracles well sclerotized and conspicuous, asperities of pronotum not very conspicuous, and paramedian anterior setae of frontal piece (frons) scarcely more caudad than the next seta, much before level of anterior sensilla; smaller seta laterad of posterior sensilla, on margin of strongly scler otized area. Conclusions from the present study differ 26 from these in that the observed pronotal asperities are quite conspicuous. Also, none of the larvae had sensilla on the frons. Aside from these differences, the observa tions of M. bicolor agreed with those traits listed by Van Emden that are unique to the species. There is much less information available on the pupae of the family Curculionidae. Burke (1968) could not determine what characters were unique to anthonomines due to the paucity of information on other weevil pupae. Information on rhynchitid pupae is also scarce, but hope fully this description along with others in progress, will provide the information necessary to define a rhyn chitid pupa. Certain characters were found to be distinct from other curculionids, and these may be characters that are unique to rhynchitids (Table 2). 27

EXPLANATION OF FIGURES 1-8

Figure Adult of M· bicolor on bud on Rosa carolina L. 2. Damage to rose bloom by adult of M. bicolor J. Oviposition-punctured rose hip 4. Egg of M· bicolor within rose hip 5. Larva of M· bicolor within rose hip 6. Larval feeding damage to rose achenes Plastic crisper for containing adults (20 em X 10 em X 8 em) 8. Plaster rearing cast for overwintering larvae (17 em X 12 em X 4 em) ,., 2u 29

EXPLANATION OF FIGURES 9-lJ

Figure • 9. James Woodworth Prairie Preserve, Niles, Illinois 10. Rose bush plastic wrap for obtaining emerging larvae 11. Excavation of prairie test plot to obtain overwintering larvae and pupae 12. Lining of prairie test plot with plastic screening 1J. Completed prairie test plot covered with hardware cloth

EXPLANATION OF FIGURES 14 AND 15

Figure 14. Larva of M· bicolor, lateral view, 19X 15. Head of M. bicolor larva, anterior view, 64X J2

Abll Ab.m:

----asp

--- -ES

,.-- -sps

"' '----a.nt

15 JJ

EXPLANATION OF FIGURES 16 AND 17

Figure 16. Head of M. bicolor larva, lateral view, 64X 17. Head of M. bicolor larva, dorsal view, 64X

• J4

-le.s

---- -Enc - --des - --pes

17 35

EXPLANATION OF FIGURES 18-20

Figure 18. Left antenna of M· bicolor larva, dorsal view, 4JX 19. Labrum of M. bicolor larva, dorsal view, 430 X 20. Labrum-epipharynx of M. bicolor larva, ventral view, 430 X -acap ,.,.,.- -- -art

___ \sl 19

- -ams -- -als

-msp -LrnR -;np

20 37 EXPLANATION OF FIGURES 21 AND 22

Figure 21. Left mandible of M. bicolor larva, dorsal view, lOOX 22. Labium of M. bicolor larva, ventral view, 4JOX 0 39

EXPLANATION OF FIGURES 23 AND 24

Figure 2). Left maxilla of M. bicolor larva, ventral view, 4JOX 24. Prothoracic spiracle of M. bicolor larva, 4JX 40

~--mxP / / /

-td 23

/ ,------Prt

-- -ATb

24 41

EXPLANATION OF FIGURES 25 AND 26

Figure 25. Thorax and abdominal segments I and II of M. bicolor larva, lateral view, 64X, indicating thoracic regions 26. Thorax and abdominal segments I and II of M. bicolor larva, lateral view, 64X, indicating chaetotaxy 42

-- -SI --- EuSt --- -Stn 25

26 4J

EXPLANATION OF FIGURE 27

Figure 27. Abdominal segments VII-X of M. bicolor larva, lateral view, 64X 44

27 45

EXPLANATION OF FIGURES 28 AND 29

Figure 28. Pupa of M. bicolor, ventral view, 16X 29. Pupa of M. bicolor, dorsal view, 16X 46 ------"''p

--sitS ---- .....,_- - -tts

28'~------PP

--....,.,---- -amP

------ItT

29 47

EXPLANATION OF FIGURE 30

Figure JO. The effect of temperature on larval emergence, 1978 (• = 1 larval emergence) 48 0 M

t-- N I• ~

!'1\ N

iii

!:'

'!! • ..... 0 ~ ((\ .r:. u. C') 0 .... 0== &.. II -II • • EXPLANATION OF FIGURE 31

Figure Jl. The effect of temperature on larval emergence, 1979 ( • = 1 larval emergence) 50 -M

••• • co

""' •••• :4i !!

1.0 0 l.t) ~ u. .c 0 0) ~ .c 0 .. II ~ • 51

EXPLANATION OF FIGURE 32

Figure J2. The effect of temperature on pupation, 1979 ( • = pupation) 52 32

30T-.r~~~.-.--.-.-.r-.--.-.-,--r-~~----r-~-r-r--r ~19 21 Z3 ZS Z1 Z9 5/1 3 5 1 'I II I~ 15 11 19 2.1 1~ Z5 2.'1 2.9 31 53 TABLE 1. SUMMARY OF SELECTED LARVAL CHARACTERS SELECT LARVAL CHARACTER RHYNCHITIDAE CURCULIONIDAE Subcutaneous present X pigment spots absent X Head retracted X X free X Antenna conspicuous X m~nute X Frontal indistinct X suture distinct X Labral rods long X short X Maxillary palp 3 X segments 2 X Anteromedian 2 X epipharyngeal setae 4-6 X Malar sensillae 13 X 9-12 X Pronotal setae 13-16 X 9-12 X Transverse 2 X abdominal folds 3 X Prodorsal setae 3-4 X 1-2 X Alar setae 2 X 1 X Spiracular 3-5 X setae 1 X Epipleural ~9 X setae 1-2 X Pedal setae 2 X 1 X Abdominal 4-6 X pleural setae 1-2 X Eusternal J X setae 2 X Spiracles bicameral X X unicameral X 54 TABLE 2. SUMMARY OF SELECTED PUPAL CHARACTERS

SELECT PUPAL CHARACTER RHYNCHITIDAE CURCULIONIDAE Distirostral 2 pairs X setae 1 pa1.r X Basirostral J pairs X setae 1-2 pa1.rs X Supraorbital absent X setae present X Anteromedian 2 pairs X setae 1 pa1.r X Anterolateral 5 pairs X setae 2-4 pairs X Posteromedian 5 pairs X setae 1-2 _pairs X Mesonotal and 4 pairs X metanotal setae 2-J pairs X absent X Laterotergal 5 pairs X setae 1-2 pairs X Sternal present X setae absent X Abdominal 6 pairs X spiracles 5 pairs X Femoral 5 X setae 0-2 X LITERATURE CITED

Anderson, D.M. 1968. Observations on the pupae of Anthonomus grandis grandis Boheman and ~· grandis thurberiae Pierce (Coleoptera:Curculionidae). Ann. Entomol. Soc. Am., 61(1):125-29. Anderson, W.H. 1947. A terminology for the anatomical characters useful in the taxonomy of weevil larvae. Proc. Entomol. Soc. Wash., 49(5):123-32. . Balduf, W.V. 1959. Obligatory and facultative insects in rose hips: their recognition and bionomics. Univ. of Illinois Press, Urbana, 194 pp. Blatchley, W.S. and C.W. Leng. 1916. Rhynchophora or weevils of northeastern America. Nature Pub. Co., Indianapolis, Ind., 683 pp. Boving, Adam G. and F.C. Craighead. 1931. An illus trated synopsis of the principal larval forms of the Order Coleoptera. Entomol. Am., 11(1-4): 1-351. Burke, H.R. 1968. Pupae of the weevil tribe Anthonomini (Coleoptera:Curculionidae). Tex. Agric. Exp. Stn. Monogr. 5, 92 PP• Buchanan, L.L. 1939. Changes of names in Carabidae and Rhynchophora. Proc. Entomol. Soc. Wash., 41: 79-82. Cassidy, James. 1888. Notes on insects and insecticides. Colo. Agric. Exp. Stn. Bull. 6, p. 18. Chittenden, F.H. 1901. Some insects injurious to the violet, rose, and other ornamental plants. U.S. Dep. Agric. Bull. No. 27, pp. 98-100. Clark, Wayne E. 1976. Notes on the life history and habits of Perigaster cretura (Herbst) (Coleoptera: Curculionidae) with descriptions of the larva and pupa. Coleopt. Bull., 30(2):159-65. • 1978. Notes on the life history and des------c-r~~-ptions of the larva and pupa of Neotylopterus pallidus (LeConte) (Coleoptera:Curculionidae), a seed predator of Forestiera acuminata (Michx.) Poir. (Oleaceae). Coleopt. Bull., 32(2):177-84. 55 56 Clark, W.E., H.R. Burke and D.M. Anderson. 1978. The genus Tychius Germar (Coleoptera:Curculionidae): larvae and pupae of some species, with evaluation of their characters in phylogenetic analysis. Proc. Entomol. Soc. Wash., 80(4):626-54. Cockerell, T.D.A. 1912. Notes and news. Entomological gleanings from all quarters of the globe. Entomol. News, 23:80-84. Cockerell, T.D.A. and R.C. Harris. 1924. A new form of Rhynchites (Coleop.:Curculionidae). Entomol. News, 35:144. Cooley, R.A. 1903. Two insect pests. Mont. Agric. Exp. Stn. Bull. 46, PP• 107-18. Crowson, R.A. 1955· The natural classification of the families of Coleoptera. Nathaniel Lloyd and Co., Ltd., London, 187 pp. Dickerson, Edgar L. 1910. Notes on Rhynchites bicolor Fabr. J. Econ. Entomol., 3:316-17. Essig, E.O. 1958. Insects and mites of western North America. Macmillan Co., New York, 1050 pp. Fabricius, Johann C. 1775. Systema Entomologiae. Lipsiae, 30 + 832 PP• 1792. Entomologia Systematica. Hafniae, Vol. 1, 330 + 538 PP• Fall, H.C. 1901. List of the Coleoptera of southern California, with notes on habits and distribution and descriptions of new species. Occas. Pap. Calif. A~ad. Sci., No. 8, p. 186. Gates, Burton N. 1909. The rose curculio (Rhynchites bicolor Fabr.) in Massachusetts. J. Econ. Entomol., 2:465-66. Gillette, C.P. and G.M. List. 1921. The western rose snout beetle, Rhl?chites wickhami Ckl. Colo. Entomol. Circ. 3 , p. 20. Green, J. Wagener. 1920. Notes on American Rhynchophora. Entomol. News, 31:193-201. Hamilton, Robert Wilson, 1969. Studies of the Rhyncho phorous families Nemonychidae, Attelabidae, and Rhynchitidae, with a revision of north American species of Attelabus Linnaeus, Rhynchites . Schneider, and Eugnamptus Schoenherr (Coleoptera: 57 Curculionoidea). Ph.D. dissertation, The Ohio State University, 492 pp.

------~~· 1971. The genus Pselaphorhynchites {Coleoptera:Rhynchitidae) in America north of Mexico. Ann. Entomol. Soc. Am., 64(5):982-96.

------~--· 1979. Taxonomic use of endophallic struc tures in some Attelabidae and Rhynchitidae of America, north of Mexico (Coleoptera:Curculionoidea), with notes on nomenclature. Ann. Entomol. Soc. Am., 72(1):29-34. Harris, Thaddeus W. 1862. A treatise on some of the insects injurious to vegetation. Crosby and Nichols, Boston, Massachusetts, 640 pp. Herbst, Johann F.W. 1797. Natursystem Aller Bekannten in -und Auslaendischen Insecten, •.. , Kaefer. Berlin, 7:1-346. Hoerner, J.L. 1936. Western rose curculio, Rhynchites bicolor wickhami Ckl. Colo. Agric. Exp. Stn. Bull. 4 32' pp. 1-19 . Kissinger, David G. 1964. Curculionidae of America north of Mexico, a key to the genera. Taxonomic Publications, Edwards Bros., Inc., South Lancaster, Massachusetts, 143 pp. Lovett, A.L. 1915. The rose curculio, Rhynchites bicolor Fab., injures blackberry buds. Rep. Dep. Entomol., Oregon Agric. Exp. Stn., pp. 150-53· Metcalf, C.L., W.P. Flint, and R.L. Metcalf. 1962. Destructive and useful insects, their habits and control. McGraw Hill Book Co., New York, 4th ed., 1087 PP• Olivier, Antoine G. 1807. Entomologie, ou Histoire Naturelle des Insects, •.• Paris, Vol. 5, 612 pp. Peterson, Alvah. 1960. Larvae of insects. Part II; Col~optera, Diptera, Neuroptera, Siphonaptera, Mecoptera, Trichoptera. Edwards Bros., Inc., Ann Arbor, Michigan, 4th ed., 416 pp.

------~· 1962. Larvae of insects. Part I; Lepidoptera and plant infesting Hymenoptera. Edwards Bros., Inc., Ann Arbor, Michigan, 4th ed., 315 pp. 58 ------=---· 1964. Entomological techniques. Edwards Bros., Inc., Ann Arbor, Michigan, lOth ed., 435 pp. Pierce, W. Dwight. 1913. Miscellaneous contributions to the knowledge of the weevils of the families Attelabidae and Brachyrhinidae. Proc. U.S. Natl. Mus.,;45(1988):365-426. Robertson, H.A. 1924. The rose curculio in Manitoba, with notes on other insects affecting roses. Annu. Rep. Entomol. Soc. Ont., 54, pp. 12-16. Rogers, C.E., W.E. Clark and H.R. Burke. 1975. Bionomics of Sibinia sulcatula (Coleoptera:Curculionidae) on mesquite in Texas. Southwest. Nat., 20(3):303-14. Schaeffer, Charles. 1905. Additions to the Coleoptera of the United States with notes on some known species. Sci. Bull. Mus. Brooklyn Inst. Arts Sci., 1(6):123-40. Schneider, David H. 1791. Anmerkung des Herausgebers. Neu. Mag. Entomol., 1:62-89. Sharp, D.E. 1889. Biologia Centrali-Americana, Insecta, Coleoptera, Curculionidae1Attelabinae. 4(3):1-40. Snodgrass, R.E. 1935. Principles of insect morphology. McGraw Hill Book Co., Inc., New York, 667 pp. Van Emden, Fritz. 1938. On the taxonomy of Rhynchophora larvae (Coleoptera). Trans. R. Entomol. Soc. Lond., 87:1-37· Wadley, F.M. 1967. Experimental statistics in entomology. Graduate School Press, U.S. Dep. Agric., Washington, D.C., 133 PP • Wangberg, James K. 1975. Biology of the thimbleberry gallmaker Diastrophus kincaidii (Hymenoptera: Cynipidae). Pan-Pac. Entomol., 51(1):39-48. Weishaupt, Clara G. 1960. Vascular plants of Ohio. Harold L. Hedrick, Columbus, Ohio, 309 pp. APPROVAL SHEET

The thesis submitted by Sheila s. Kuritsky has been read and approved by the following committee: Dr. Robert w. Hamilton, Director Associate Professor, Biology, Loyola Dr. Benedict J. Jaskoski Professor, Biology, Loyola

Dr. John J. Peluso Assistant Professor, Biology, Loyola The final copies have been examined by the director of the thesis and the signature which appears below verifies the fact that any necessary changes have been incorporated and that the thesis is now given final approval by the Committee with reference to content and form. The thesis is therefore accepted in partial fulfillment of the requirements for the degree of Master of Science.

Xi/ -;o-!? 71 Date