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University Microfiims INFORMATION TO USERS This dissertation was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or. chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginrimg below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. University Microfiims 300 North Zeeb Road Ann Arbor, Michigan 48106 A Xerox Education Company 72-29,881 GREEN, Edwin Alfred, 1931- THE COMPARATIVE LARVAL MORPHOLOGY OF REPRESENTATIVE NEW WORLD DRYOPOID BEETLES. The University of Oklahoma, Ph.D., 1972 Zoology University Microfilms, A XEROX Company, Ann Arbor, Michigan THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED THE UNIVERSITY OF OKLAHOMA nPAOUATE COLLEGE THE COMPARATIVE LARVAL MORPHOLOGY OF REPRESENTATIVE NEW WORLD DRYOPOID BEETLES A DISSERTATION SUBMITTED TO THE GRADUATE FACULTY In partial fulfillment of the requirements for the degree of doctor of philosophy BY EDWIN ALFRED GREEN Merman, Oklahoma 1972 THE COMPARATIVE LARVAL MORPHOLOGY OF REPRESENTATIVE NEW WORLD DRYOPOID BEETLES APPROVED BY DISSERTATION C0f4MITTEE PLEASE NOTE: Some pages may have indistinct print. Filmed as received. University Microfilms, A Xerox Education Company ACKNOWLEDGMENTS The w rite r wishes to express his appreciation and thanks to Dr. Harley P. Brown for his guiding assistance and criticism during the course of th is study and the preparation of the dissertation. Dr. Brown also kindly contributed most of the larvae studied here and made available to me reference materials without which this study could not have been completed. I also wish to thank Drs, George Goodman, Cluff Hop la, and Frank Sonleltner for their assistance and criticism of this study. I am deeply obliged to David Shetlar for his assistance on a collecting trip to southern Oklahoma. Finally, I would like to thank my wife, Loretta for her continued encouragement and Pamela, Edwin J r ., and Eric fo r th e ir patience. III TABLE OF CONTENTS Page LIST OF FIGURES v I. INTRODUCTION 1 I I . MATERIALS AND METHODS 3 I I I . RESULTS 7 PsephenIdae 7 Llmnlchldae 54 Elmldae 84 IV. DISCUSSION 376 V. SUMMARY AND CONCLUSIONS 387 LITERATURE CITED 389 Iv LIST OF FIGURES Page(s) External Morphology of Psephenus texanus (Figures 1-7) 32-36 Internal Morphology of Psephenus texanus (Figures 8-10) 37-38 Histological Morphology of Psephenus texanus (Figure 11) 39 External Morphology of Ectoprla nervosa (Figures 12-19) 40-44 Internal Morphology of Ectoprla nervosa (Figures 20 - 22) 45 Histological Morphology of Ectoprla nervosa (Figures 23 - 26) 46-48 External Morphology of Psephenops smith I (Figures 27 - 33) 49-50 Internal Morphology of Psephenops smith! (Figures 34 - 35) 51 Histological Morphology of Psephenops smith! (Figures 36 - 38) 52-53 External Morphology of Lutrochus luteus (Figures 39 - 48) 69-72 Internal Morphology of Lutrochus luteus (Figures 49 - 50) 73 Histological Morphology of Lutrochus luteus (Figures 51 - 53) 74-75 External Morphology o f Lutrochus "8” (Figures 54 - 68) 76-81 Internal Morphology of Lutrochus "B” (Figures 69 - 71) 82 Histological Morphology of Lutrochus "B" (Figures 72 - 73) 83 External Morphology of Hexanchorus caralbus (Figures 74 - 87) 207-210 Internal Morphology of Hexanchorus caralbus (Figures 88 - 90) 211 Histological Morphology of Hexanchorus caralbus (Figure 91) 212 External Morphology of Lara avara (Figure 92 - 102) 213-219 Internal Morphology of Lara avara (Figure 103) 220 Histological Morphology of Lara avara (Figures 104 - 108) 221-222 External Morphology of PotamophIlops (Figures 109 - 123) 223-229 Internal Morphology of PotamophItops (Figures 124 - 125) 230 Histological Morphology of PotamophIlops (Figures 126 - 129) 231 External Morphology of Phanocerus clavlcornis (Figures 130 - 142) 232-235 Internal Morphology of Phanocerus clavlcornis (Figures 143 - 144) 236 Histological Morphology of Phanocerus clavlcornis (Figures 145 - 147) 237 External Morphology of Ancyronyx varlegata (Figures 148 - 163) 238-247 Internal Morphology of Ancyronyx varlegata (Figures 164 - 165) 248 Histological Morohology of Ancyronyx varlegata (Figures 166 - 169) 249-251 vl External Morpholoqy of Austrolimnîus formosus (Figures 170 - 187) 252-256 Histological Morphology of Austrolimnîus formosus (Figures 189 - 191) 257 External Morphology of EIslanus texanus (Figures 192 - 208) 258-266 Internai ?4orpholoqy of EIslanus texanus (Figuras 209 - 210Ï 267 Histological Morphology of EIslanus texanus (Figures 211 - 218) 268-271 External Morphology of Macronychus qîabratus (Figures 219 - 240) 272-281 Internai Morphology of Macronychus qîabratus (Figures 241 - 242) 282 Histological Morphology of Macronychus olabratus (Figures 243 - 248) 283-284 External Morphology of Neocylloepus sandersoni (Figures 249 - 262) 286-291 Histological Morphology of Heocylloepus sandersoni (Figures 263 - 265) 2.92-294 External Morphology of Nooeimis sp» (Figures 266 - 276) 295-299 Histological Morphology of Meoelmls sp. (Figure 277) 300 External Morphology of Optlosorvus oval Is (?) (Figures 278 - 237) 301-307 Internal Morphology of Optloservus oval Is (?) (Figures 288 - 289) 308 Histological Morphology of Optloservus oval Is (?) (Figures 290 - 292) 309 External Morphology of Oullmnlus iatlusculus (Figures 293 - 302) 310-314 Histological Morphology of Oullmnlus iatlusculus (Figures 303 - 304) 315 External Morphology of Promoresla tarde 1 la (Figures 305 - 316) 316-320 vil Hîstologfcal Morphology of Promorasla fardel la (Figures 317 - 318) “ 321-321 External Morphology of Tolrlolus ungulatus (Figures 319 - 331) 323-328 Internal Morphology of Tolrlolus ungulatus (Figure 332) ' 329 Histological Morphology of Tolrlolus ungulatus (Figure 333) 330 External Mofphology of Zaltzovla parvula (Figures 334 - 3481--------------------------- 331-337 Internal Morphology' of Zaltzavla parvula (Figures 349 - 350')’ “"“" 338 Histological Morphology of Zaltzovla parvula (Figure 351) 339 External Morphology of Xonelmls bufo (Figures 352 - 365)-------------------- 340-344 Internal Morphology of Xenelmis bufo (Figures 366 - 367)--------------------- 345 Histological Morphology of Xenelmis bufo (Figure 368) 346 External Morphology of MIcrocylloepus pustllus (Figures 369 - 3807 ---------------- 347-353 Internal Morphology of MIcrocyl loepus pustllus (Figure 381) 354 Histological Morphology of MIcrocylloepus puslllus (Figura 352) 355 Exfornal Morphology of Dub I raphia sp. (Figures 383 - 394)' ................. 356-359 Internal Morphologyof Dublraphia sp. (Figures 395 - 391Î) -------------- 360 External Morphology of Heterelmls vulnerata (Figures 397 - 408) --------------------------------- 361-366 Histological Morphology of Heterelmls vulnerata (Figures 409 - 4 1 2 ) ---------------------------------- 367-371 vlll External Morphology of Stenetmis sp. (Figure 413 - 420) 372-374 Internal Morphology of Stenelmis sp. (Figure 421) 375 ix THE COMPARATIVE LARVAL MORPHOLOGY OF REPRESENTATIVE NEW WORLD DRYOPOlD BEETLES INTRODUCTION With the publication of Leng's checklist (1920), taxonomists be­ came aware of the need fo r a more natural system of classifying the Coleoptera, Such a system could only be developed when an adequate consideration of larval features Is taken Into account along with adult features. In spite of general recognition of the need for studies of larvae, our present knowledge of the morphology of Immature dryopold beetles results from the work of a very limited number of Investigators. The classical work by Bbvlng and Craighead (1931) represents the largest single publication concerned with the i vae of beetles. This work, which Included a number of dryopolds, caused Lang and Mutch 1er (1933) to make sig n ifican t changes In th e ir subsequent classificatio n scheme. West (1929), prsssr/lng the family name of Dryopldae, studied the external morphology of larval structures In selected genera. Later studies by other Investigators have necessitated changes In most of the names used by West, but his work represented a great contribution to our
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