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| University Microfilms, a Xeroxcompany, Ann Arbor, Michigan 72-4557 MANISCHEWITZ, Jack Roger, 1942- - • A NUMERICAL PHENETIC STUDY OF THE SNAKE MITES OF THE FAMILY IXOD ORHYNCHIDAE (ACARI: MES0STI6MATA). The Ohio State University, Ph.D., 1971 Entomology | University Microfilms, A XEROX Company, Ann Arbor, Michigan — a . •*- ' T THTR T1TRRERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED A NUMERICAL PHENETIC STUDY OF THE SNAKE MITES OF THE FAMILY IXODORHYNCHIDAE (ACARI: MESOSTIGMATA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Jack Roger Manischewitz, B.A., M.S ****** The Ohio State University 1971 Approved by Adviser Department of Entomology PLEASE NOTE: Some Pages have Indistinct print. Filmed as received. UNIVERSITY MICROFILMS ACKNOWLEDGMENTS i The following individuals and institutions graciously provided the specimens used in the present study: J. Camin (University of Kansas), A. Fain (Institut de Medecine Tropicale, Antwerp, Belgium), 2. Feider (Universitatea "Al. I. Cuza," Jasi, Romania), W, Voss (Fort Worth Children*s Museum), The Acarology Laboratory (Columbus, Ohio), J. Cooreman (Institut Royal des Sciences Naturelles de Belgique, Brussels), and M. Naudo (Museum National D'histoire Naturelie, Paris). Drs. F. Rohlf and J. Kishpaugh (SUNY, Stony Brook) generously provided.the computer program NT-SYS. Thomas Kozlowski provided aid in the use of computers. Free computer time was donated by the Instructional and Research Computer Center of The Ohio State University. I would like to acknowledge gratefully the important guidance and assistance provided by my adviser, Dr. Donald Johnston. This study was supported in part by a predoctoral traineeship from the National Institutes of Health Training Grant 5 T01 A100216. ii VITA i March 31, 1942. Born - Cincinnati, Ohio 1964................ B.A., Rutgers— The State University, New Brunswick, New Jersey 1964-1965 .......... Research Assistant, Department of Entomology and Economic Zoology, Rutgers— The State University, New Brunswick, New Jersey 1966............... M.S., Rutgers— The State University, New Brunswick, New Jersey 1966-1969 .......... Teaching Assistant, The Ohio State University, Columbus, Ohio 1969-1971 .......... Acarology Predoctoral Trainee, Acarology Laboratory, The Ohio State University, Columbus, Ohio PUBLICATION Manischewitz, Jack. 1966. Studies on Parasitic Mites of New Jersey. Jour. New York Ent. Soc. 74:189-97. FIELDS OF STUDY Major Fields: Zoology and Entomology Studies in Acarology and Numerical Taxonomy. Professors D. E. Johnston and 6 . W. Wharton iii TABLE OF CONTENTS i ACKNOWLEDGMENTS................................................ ii VITA.............................................................. Ill LIST OF TABLES.................................................. v LIST OF ILLUSTRATIONS.......................................... Vi INTRODUCTION................................................... 1 MATERIALS AND METHODS.......................................... 4 RESULTS AND DISCUSSION................. 15 Predictiveness Determination and a Consideration of Weighting............. 15 The Ixodorhynchidae .................. ......... 31 SUMMARY....................................................... 53 APPENDIX I. LIST OF O T U ' S ..................................... 55 II. CHARACTERS U S E D .................................... 62 III. BASIC DATA MATRIX FOR 103 SPECIMENS............ 67 FIGURES.......................................................... 112 REFERENCES ........... 120 iv LIST OF TABLES Table 1. List of Included Species and Their Code Names. ...... 5 2. Summarization of Characters........... ... 7 3. Stability Predictiveness and Character Predictiveness of Similarity Matrices............... 24 4. Stability Predictiveness and Character Predictiveness of Rienograms........... ................................. 25 v LIST OP ILLUSTRATIONS Figure 1. Standardization M.C.D.. Phenogram........................ 113 2. Graph Analysis of Ixodorhynchus. Ixobioides. and Ixodorhvnchoides, Based on the Standardization M.C.D. Similarity Matrix................................ 115 3. Graph Analysis of Hemilaelaps, Scutanolaelaps. Strandtibbettsia . and Omentolaelaps. Based on the Standardization M.C.D. Similarity Matrix, ...... 117 4. Graph Analysis of Nearest Neighbors of Hemilaelaps triangulus, Based on the Standardization M.C.D. Similarity Matrix ............... 119 vi INTRODUCTION The present study examines relationships within the family Ixodorhynchidae Ewing, 1923, using the principles and techniques of numerical taxonomy. In addition, an attempt is made to provide infor­ mation relevant to numerical taxonomy in general. A variety of transformations and similarity coefficients were used in producing a variety of similarity matrices and phenograms (Sokal and Sneath, 1963). The techniques of principal components (Harman, 1967) and graph analysis (Moss, 1967) were also used. Prediction is often regarded as the major goal of classification (Gilmour, 1940; Inglis, 1970; Sokal and Sneath; 1963). However, there has been little attempt to determine empirically the relative predictive success of similarity matrices or phenograms produced by alternative methods. Such an attempt is made in this study, primarily on the basis of randomly dividing characters into groups and finding the correlations between various matrices and phenograms. As a result of a consideration of the meaning of character weighting, a new transformation, equalization, is proposed. It is designed to give characters equal weight. In most numerical taxonomic studies, OTU's are species or other groupings of individuals. However, an alternative method, in which individual specimens are the OTU's, is also used (e.g., Moss, 1967, and Herrin, 1969). The present study used both methods. 1 Ixodorhynchids are gamasine mites ectoparasitic on snakes.. Radovsky (1969) believes the Ixodorhynchidae arose from the Laelapinae, and gave rise to the Omentolaelapidae and Eiitonyssidae. Fain (1962) reviewed the Ixodorhynchidae, and readers interested in the history of the family prior to 1962 should consult this work. Fain's study was based on conventional taxonomic methods. Taxa were determined on the basis of few characters, primarily those of chelicerae, cornlculi, and coxal setae. He recognized the following genera: Ixodorhynchus Ewing, 1923, Ixobioides Fonseca, 1934, Hemilaelaps Ewing 1933, Asiatolaelaps Fain 1961, and Strandtibbettsla Fain, 1961. He further divided Hemilaelaps into 4 "groups" on the basis of coxal setae: tri'angulus group, farrieri group, piger group, and ophidius group, the . last corresponding to the genus Scutanolaelaps Lavoipierre, 1958, sunk by Fain on the basis that one species, £». upembae Fain, 1961, was inter- l mediate between the two genera. In 1962, • Johnston described Ixodorhynchus neodelphus. I, faini, and the new genus and species Ixodorhynchoides truncatus. Voss and Strandtmann (1962) described Ixodorhynchus uncatissimus, which, according to Fain's criteria, would be placed in Ixobioides. Taufflieb (1967), unaware of the Fain (1962) paper sinking Scutanolaelaps. described Scutanolaelaps mehelyae. Voss (1967) described Hemilaelaps phillippinensis. which because he believed it to be intermediate between Hemilaelaps and Asistolaelaps. caused him to synonymize Asiatolaelaps with Hemilaelaps. H. lioheterodon was described by Fain (1967). J. H. Gamin, University of Kansas, has discovered a new species, which, according to Fain's (1962) criteria, would belong in Strandtibbettsia; however, Gamin has suggested that it represents a new genus. The history of the family has been marked by the sole use of traditional taxonomic methods, including use of few characters in determining taxa. Due in part- to the emergence of papers during the same year (3 in 1962 and 3 in 1967), workers have sometimes been unaware of each other's work. Further, a preliminary examination of some characters not considered by Fain suggested that some generic changes would be appropriate, particularly with regard to Ixodorhynchus, Ixobioides. Strandtibbettsia. and Asiatolaelaps. The above factors have contributed to the present state of generic uncertainty, which hopefully can be ameliorated by the present study. MATERIALS AND METHODS i A total of 103 specimens, representing 33 taxonomic species were included in the study. All were adult females, as immature stages and males, were not generally available. Additional available specimens were omitted on the grounds that they had too much missing data, or that the species involved were already well-represented. Within each species an attempt was made to use specimens with differing host-locality data. The number of specimens used per species varied from 22 to 1. Of the 32 currently recognized species of Ixodorhynchidae, all were available ' for inclusion except Hemilaelaps imphalensis (Radford, 1947). Also included were representatives of a new species found by Camin, and Omentolaelaps mehelyae Fain, 1961, the single member of the family Omentolaelapidae. Table 1 lists the species used and gives the code names used in the phenograms and graph analyses. Appendix I lists the specimens used, and gives host-locality data. Specimens were observed by means of a phase contrast microscope (American Optical). A micrometer disc in the eyepiece was used to obtain measurements. A total of 190 characters were originally considered, and the states of these 190 characters in each of the 103 specimens were recorded. For
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