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Phylogenetic Relationships in Neotomine-Peromyscine Rodents (Muroidea) and a Reappraisal of the Dichotomy Within New World Cricetinae

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Phylogenetic Relationships in Neotomine-Peromyscine Rodents (Muroidea) and a Reappraisal of the Dichotomy Within New World Cricetinae MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 157 Phylogenetic Relationships in Neotomine-Peromyscine Rodents (Muroidea) and a Reappraisal of the Dichotomy within New World Cricetinae by Michael Dean Carleton National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN December 12, 1980 MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN The publications of the Museum of Zoology, University of Michigan, consist of two series - the Occasional Papers and the Miscellaneous Publications. Both series were founded by Dr. Bryant Walker, Mr. Bradshaw H. Swales, and Dr. W. W. Newcomb. The Occasional Papers, publication of which was begun in 1913, serve as a medium for original studies based principally upon the collections in the Museum. They are issued separately. When a sufficient number bf pages has been printed to make a volume, a title page, table of contents, and an index are supplied to libraries and individuals on the mail- ing list for the series. The Miscellaneous Publications. which include .DaDers . on field and museum tech- niques, monographic studies, and other contributions not within the scope of the Occa- sional Papers, are published separately. It is not intended that they be grouped into volumes. Each number has a title page and, when necessary, a table of contents. A complete list of publications on Birds, Fishes, Insects, Mammals, Mollusks, and Reptiles and Amphibians is available. Address inquiries to the Director, Museum of Zool- ogy, Ann Arbor, Michigan 48109. MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN NO. 157 Phylogenetic Relationships in Neotomine-Peromyscine Rodents (Muroidea) and a Reappraisal of the Dichotomy within New World Cricetinae hy Michael Dean Carleton National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 Ann Arbor MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHlGAN December 12, 1980 CONTENTS PAGE INTRODUCTION ............................ .............................. 1 Background of the Problem ..................... .............................. 2 Objcctivcs of the Study ........................ ACKNOWLEDGMENTS ....................... MATERIALS AND METHODS ................. Selection of Species' Samples .................... Character Description and Numerical Methods .... SURVEY OF CHARACTERS ................... Dentition .................................... Cranium ..................................... Post-Cranial Skeleton .......................... Alimentary Canal ............................. Phallus ...................................... Male Accessory Reproductive Glands ............. Miscellancous External Features ................. RESULTS ..................................... Phenetic Analyses of Neotomine-Peromyscines ..... Phylogenetic Analyses of Neotomine-Peromyscines ................................... 90 Morphological Variation of the Glans Pcnis in New World Cricetines andOtherMuroidea .......................................................... 105 Neotomine.Peromyscines. South American Cricctincs. andotherGroupsofMuroidea .................................................. 110 DISCUSSION ................................................................... 113 Relationships Within Neotomine-Peromyscines ...................................... 113 The Simple Versus Complex Penis and a Reappraisal of the Neotomine-Peromyscine and South American Cricetine Dichotomy ................... 127 LITERATURECITED ........................................................... 140 ILLUSTRATIONS FIGURE PAGE 1. Hierarchical arrangement of major groups of Muroidea .............................. 7 2 . Collective distribution of South American cricetincs ................................. 8 3 . Collective distribution of neotomine-peromyscines .................................. 9 4 . Relationships of neotomine-peromyscines as proposed by HooperandMusscr(1964b) .................................................... 11 5 . Frequency histograms used to compute Character Nos . 1,6,33. and 42 ................. 38 6 . Alveolar patterns of upper and lower cheek teeth .................................... 40 7 . Schematic diagram of carotid circulatory patterns (ventral aspect) illustrating transitional states dcfined for Character No . 16 ........................... 43 8 . Ventral view of rnesopterygoid fossa showing absence and presence ofsphenopalatinevacuities ..................................................... 45 9 . Variation in posterior palatine foramina .......................................... 46 10 . Ventral view of lower jaws illustrating posteriorly directed angular process and deflected angular process ...................................... 48 11. Ventral view of the hyoid bones of Neolomafusclpes and Sigmodon hispidus ................. 49 12 . Left mallei of Neolomu lepida and Dicroslonyxgroenlandicus .............................. 50 13. Distal portion of left humeri demonstrating presence and absence ofentepicondylarforamcn ...................................................... 53 14. Frontal aspect of second thoracic vertebrae, one having pronounccd neural spine and the other lacking this process ...................................... 54 15. Dorsal view of seventh cervical and first thoracic vertehrac, demonstrating articular arrangements of the first rib ................................ 55 16. Left calcanea of Neotoma cinerea, Holochilus brasiliensis and 7hteru indica ................... 56 17 . Left scapulae of Peromyscus boylii and Gerbillw cheesmani illustrating development of third scapular fossa ..................................... 56 18. Differences in degrce of fusion of the tibia and fibula as seen in Tylomys nudicaudus and Reilhrodonlomys creper ........................................ 57 19. Schematic diagrams of complex and simple glandes penis as secn from lateral view ........ 60 20 . Frequency histograms used to compute proportional characters of the glans penis ......... 63 21 . Ventral view of male reproductive tract of Nectomys squam$es illustrating proposed ancestral complement of accessory glands ................................. 66 22 . Scatter plot of principal component I versus I1 (A) and I versus 111 (B) using 47 species of neotomine-perornyscines as dcscribed by 60 characters .................... 78 23 . Cluster analysis (unweighted, pair-group method using arithmetic averages) of 47 ncotominc-peromyscines based on principal components one through ten ............. 80 24 . Scatter plot of principal component I versus I1 (A) and I1 versus 111 (B) using 37 species of Peromyscini as described by 52 characters .............................. 86 25 . Scatter plot of principal component I versus I1 using 31 species of Peromyscus and related genera as described by 48 characters .................................... 88 26 . Scatter plot of principal component I versus I1 (A) and I versus I11 (B) using 11 species of Neotomini and Nelsonia as described by 31 characters ..................... 90 27 . Shortest connection network (Prim) of44 species of neotomine-peromyscines ............ 92 28 . Wagner tree analysis of 44 species of neotomine-peromyscines based on 59characters ................................................................. 94 FIGURE PAGE 29. Wagner tree analysis of 34 species of neotomine-peromyscines based on 59 characters . 95 30. Wagner tree analysis using hypothesized common ancestors of certain taxa . 98 31. Weighted Wagner tree analysis of 44 species of neotomine-peromyscines . 100 32. Tree produced by weighted invariant step strategy . 101 33. Estimate of phylogeny obtained with character compatibility analysis of 34 species of neotomine-peromyscines . 103 34. Estimate of phylogeny obtained with character compatibility analysis of 9 species of Neotoma and Nelsonia neotomodon . 104 35. Scatter plot of principal component I versus I1 (A) and I versus I11 (B) using all 75 species as described by 16 characters of the glans penis . 106 36. Scatter plot of principal component I versus I1 and shortest-connection network overlay, using 71 species as described by 16 characters of the glans penis . 109 37. Wagner tree analysis of all 75 species based only on the 16 characters of the glans penis and assuming the "complex" type is ancestral . 11 1 38. Wagner tree analysis of all 75 species based only on the 16 characters of the glans penis and assuming the "simple" type is ancestral. 112 39. Scatter plot of principal component I versus I1 (A) and I versus I11 (B) using 69 species as described by 74 characters . 114 40. Shortest connection network (Prim) of 56 species representing neotomine- peromyscines, South American cricetines and Old World cricetines . 116 41. Wagner tree analysis of 58 species representing neotomine-peromyscines, South American cricetines, and Old World cricetines based on 62 characters . 117 42. The phylogeny of neotomine-peromyscine rodents proposed herein . ,118 43. Character compatibility of the glans penis with five other anatomical featuresfortaxaofMuroidea .................................................... 134 44. Summary of principal trends in evolution of the glans penis based on my interpretation of character polarities and superimposed on the results of principal component analysis . 136 LIST OF TABLES TABLE PAGE 1. Number of living species of Muroidea in relation to total number ofMammalia ................................................................ 1 2. Previous
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