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Universiv Miaxsilms International INFORMATION TO USERS This was produced from a copy of a document sent to us for microfilming. 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 material submitted. The following explanation of techniques is provided to help you understand markings or notations which may appear on this reproduction. 1. The sign or “ target” for pages apparently lacking from the document photographed is “Missing Page(s)”. I f it was possible to obtain the missing page(s) or section, they are spliced into the Him along with adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure you of complete continuity. 2. When an image on the Him is obliterated with a round black mark it is an indication that the Him inspector noticed either blurred copy because of movement during exposure, or duplicate copy. 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ZEEB ROAD, ANN ARBOR, Ml 48106 18 BEDFORD ROW, LONDON WC1 R 4EJ, ENGLAND 8027532 SHEPARD, WILLIAM DOUGLAS A MORPHOLOGICAL STUDY OF LARVAL FORMS OF NORTH AMERICAN SPECIES OF STENEIMIS (COLEOPTERAzELMIDAE) The IMiversity of Oklahoma PH.D. 1980 University Microfilms I ntern ationel 300 N. Zeeb Road, am Arbor, M I 48106 THE UNIVERSITY OF OKLAHOMA GRADUATE COLLEGE A MORPHOLOGICAL STUDY OF LARVAL FORMS OF NORTH AMERICAN SPECIES OF STENELMIS (COLEOPTERA:EIMIDAE) A DISSERTATION SUBMITTED TO THE GRADUATE FACULTY in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY BY WILLIAM DOUGLAS SHEPARD Norman, Oklahoma 1980 A MORPHOLOGICAL STUDY OF LARVAL FORMS OF NORTH AMERICAN SPECIES OF STENELMIS (COLEOPTERA:EIMIDAE) APPROVED BY DISSERTATION COMMITTEE ACKNOWLEDGEMENTS A special debt of thanks is owed to Dr. Harley Brown. He suggested this problem, frequently offered help and suggestions and critically reviewed this manuscript. Additionally, he provided most of the specimens used in the analysis. Further, his continual support and interest in this student, as with many others, has enhanced my general education, attitudes and perspectives. Appreciation is also extended to Drs. Cluff Hopla, James Robertson, Gary Schnell and Frank Sonleitner. In addition to serving as my dissertation committee, all have freely offered advice concerning many subjects and have also enhanced my general education. For the loan of specimens and/or for the sharing of information relative to Stenelmis life histories, I wish to thank the following people: Drs. David White, Henry Seagle and Randal Farmer. For assistance in computer analysis, I wish to thank Mr. James Felly, Mr. Anthony Schene, Mr. William Magdych and, especially. Dr. Michael Douglas. iii tab le of contents Page LIST OF FIGURES V LIST OF TABLES vi I. INTRODUCTION 1 Stenelmis in North America 1 Other elmld larval research 2 Need for research 5 Life cycle 6 II. METHODS AND MATERIALS 13 III. GENERAL MORPHOLOGY 16 Setatlon and surface ornamentations 16 External anatomy of the larva of Stenelmis 20 IV. SYSTEMATIC TREATMENT 37 Key to the species 38 Species descriptions 41 V. DISCUSSION 69 LITERATURE CITED 87 APPENDIX 119 Iv LIST OF FIGURES Pages Setatlon and Surface Ornamentations 19 (Figures I-11) Correlation Phenogram of the 32 Characters used In the 73 Numerical Analysis (Figure 12) Distance Phenogram of the Larvae of 14 Species of Stenelmis 80 (Figure 13) Stick and Ball Model of the Distribution of Species in the 82 Space Described by the First Three Components from the Principal Components Analysis (Figure 14) Wagner Network Illustrating the Relatedness of 14 Species 85 of Stenelmis (Figure 15) External Anatoay of the Mature Larva of Stenelmis beameri 94-118 (Figures 16-51) Variations In the External Anatomy of Mature Larvae of 123-127 Stenelmis (Figures 52-73) LIST OF TABLES Pages 1, Taxonomic Actions Involving North American Species of 3 Stenelmis 2, Works Involving Larvae of North American Species of 4 Stenelmis 3, Characters Used to Evaluate the Species Numerically 70 4, Means and Standard Deviations for Characters Used 7 % in the Numerical Analysis 5, Correlation of Means of Counts and Pronotal Length for 75 all Species 6 , Correlation of Means of Counts and Pronotal Length for 7 5 Stenelmis beameri 7, Character Loadings on the First Three Principal 7 7 Components V i A MORPHOLOGICAL STUDY OF LARVAL FORMS OF NORTH AMERICAN SPECIES OF STENELMIS (COLEOPTERA:ELMIDAE) INTRODUCTION Stenelmis in North America The genus Stenelmis was erected in 1835 by Leon Dufour for the European species Elmis canaliculate. Shortly thereafter, the North American species Elmis crenata Say was reassigned to Stenelmis. Between 1852 and 1938, 12 more species of Stenelmis were described from this continent. In 1938, M. W. Sanderson published a monograph on North American species of Stenelmis in which 18 new species were described and four previously recognized species were relegated to synonymy. Sanderson's monograph has remained the largest single work on North American species of Stenelmis. Since 1938, only two new species have been described although approximately six recognized but undescribed species are known to exist. Table 1 summarizes additions and deletions to the list of North American species. In North America, very little has been done with larvae except in connection with characterizing the family Elmidae (BUving and Craighead 1930, Peterson 1951) or characterizing genera within the family (West 1929, Sanderson 1938 and 1954, Green 1972, Brown 1972). Larval descriptions exist for two species of Stenelmis— S^. bicarinata (Matheson 1914) and crenata (Boving and Craighead 1930, LeSage and Harper 1976a and 1977)— although they are generally too brief to help distinguish the species. In addition to these studies, the larval respiratory system of S. sulcatus (=S^. quadrimaculatus) was described in detail by Susskind (1936) and the life cycle has been worked out for 2" sexlineata (White 1978), S^. crenata (LeSage and Harper 1978, Seagle 1978) and S_. exigua (Farmer 1978). Table 2 gives the chronology of these developments. Worldwide, much of the work involving larval Stenelmis and other elmids has been done by H. Bertrand (1939, 1967, 1969 and 1972). In England canaliculata has been studied by Holland (1972) and Claridge and Staddon (1960). This same species was studied in Italy by Saraceni (1969). Other Elmid Larval Research The larvae of several other genera of elmids have been investigated morphologically. They include; Elmis (Berthelemy and Riols 1965), Limnius (Berthelemy 1967, Berthelemy et Stragiotti 1965), Riolus (Marvillet 1960, Berthelemy et Stragiotti 1965), Macronychus (Perez 1863), Esolus and Oulimnius (Berthelemy et Ductor 1965), Normandie and Grouvellinus (Bertrand 1967) and Zaitzevia (Pennak 1953). In addition, larvae of many genera have been treated morphologically and illustrated in a number of works dealing with elmids from a restricted area (Bertrand 1939, 1969, 1972, 1973 and 1974; Holland 1972; Brown 1972). The structure and function of the anterior digestive tract in elmid larvae has been studied by Delachambre (1964). Finally, Hinton (1939) and Green (1972) used larval morphology as part of the basis for revising the higher classification in the superfamily Dryopoidea. Table 1. Taxonomie actions involving North American apeclea of Stenelmia. Date Author Taxa described Chanaes 1824 Say Elmis crenata 1835 DuFour Stenelmis 1852 LeConte S. sinuata. S. bicarinata crenata reassigned to Stenelmis 1859 . Mbtschulsl^ S. humerosa. S. markelii. S. sordida. S. eloneata 1869 Zimmerman S. linearis. S. vittipennis 1870 Horn S. quadrimaculata S. linearis " S. humerosa! 1901 Fall S. nubifera 1910 Blatchley s. sulcata 1925 Blatchley s. fuscata w 1933 Husgrave sulcata preoccupied. renasMd blatchlevi 1938 Sanderson s. concinna. S. tarsaiis. S. knobeli S. sordida “ S. crenata s. exilis. S. mera. S. douglasensis • S. blatchlevi " S. auadrimaculata s. mirabilis. S. sexlineata. S. antennalia S. elonaata - S. canaliculata (not S. exlRua. S. beameri. S, lateralis in N. America) s. museravei. S. decorata. S. convexula - 1940 Musgrave S. tarsaiis preoccupied, renamed S. sandersoni 1949 Chandler s. calida 1958 Sanderson S. nubifera reassigned to Ordobrevia 1978 White and s. Rammoni Brown Table 2, Works Involving larvae of North American species of Stenelmis. Date Author Contents 1914 Matheson Describes S. bicarinata larva and pupa 1929 West Describes larval morphology of Stenelmis 1930 Boving and Illustrates larval structures of S. crenata Craighead 1936 Susskind Describes larval respiratory system for S. quadrlmaculata
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