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Mammalian Taxonomy and Community Ecology of a Neotropical Mountain Range

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Mammalian Taxonomy and Community Ecology of a Neotropical Mountain Range ISLANDS IN THE SKY: MAMMALIAN TAXONOMY AND COMMUNITY ECOLOGY OF A NEOTROPICAL MOUNTAIN RANGE by KATHLEEN BELVIN BLAIR Bachelor of Science Kansas State University Manhattan, Kansas 1975 Master of Science Oklahoma State University Stillwater, Oklahoma 1978 Submitted to the Faculty of the Graduate College of the Oklahoma State University in partial fulfillment of the requirements for the Degree of DOCTOR OF PHILOSOPHY JULY, 1989 ' • ...,, ·.I ' '). Oklahoma State Univ. Librar ISLANDS IN THE SKY: MAMMALIAN TAXONOMY AND COMMUNITY ECOLOGY OF A NEOTROPICAL MOUNTAIN RANGE Thesis Approved: ______ ,.,. C 0 P Y R I G H T by Kathleen Belvin Blair July 29, 1989 PREFACE Tropical ecosystems are among the most endangered and least known on earth even though they contain at least 50% of the world's species. Tropical mountains exhibit a rich diversity of terrestrial habitats produced by altitudinal changes affecting temperature and soil moisture which serve to compress many habitats into relatively small geographical areas. In addition, tropical mountains have acted as refugia for species during changes in the paleoclimate. The diversity of habitats combined with refugial effects and the species packing common to tropical systems should result in communities of unusual complexity and interest in neotropical mountain ranges. Data were collected from May 1979 - October 1980 in the Serra do Capara6 mountains of eastern Brazil" (lat. 2f 22' to 0 20 40', Lon. 40'W. to 42'W.) and were used to describe mammalian communities present and the effects of altitude and season on them. MacArthur's (1972) hypothesis predicting lower .species diversity on tropical mountaintops relative to their lower elevations when compared to temperate mountains is tested. Results of this study are discussed in light of island biogeography (MacArthur and Wilson 1967), and refugia as a force in tropical speciation (Hafter 1979; Brown 1982) is considered. Taxonomic analyses of Caparao rodents ar~ presented along with life history notes for all species found in the park. This dissertation evolved out of a project begun while I was a Peace Corps Volunteer assigned to the Instituto Brasileiro do Desenvolvimento Florestal, Minas Gerais, Brazil. It was ori6inally through the foresight and efforts of the Peace Corps, Dr. Tracy S. Carter and particularly Dr. Ivens Pinto Franyueira that this project was created to survey the mammalian fauna of Caparao and other national parks in Brazil with the goal of developing management plans for those few, precious areas. Dr. Iven's determination and dedication to the conservation and wise use of natural areas and resources and his deep concern for the preservation of endangered species and their habitats should be a model for future generations of conservationists in Brazil and throuJhout the world. IBDF generously supplied equipment, quarters, transportation and all possible support. In µarticular the .assistance of Gloria de Fatima dos Santos Mindes, biolo8ist for IBDF/MG is gratefully acknowledged. The personnel of Caparao National Park itself, both administrators and most especially the park rangers, were enormously helpful and without their day-to-day co-operation and willing hard work, little of this would have been , / µossible. E tambiem, a gente boa do Alto Caparao - Jovericy, Donna Maria, Macenieas, Hamilton e muitos otros, sem vo~es, / sem sua amazade e espiran~as, nada disso ficaria possivel. , Muitas obrigatas tem que ir aos todos de VOfeS. iv Only through the concern and efforts of many people has this dissertation come about. My committee, Drs. James H. Shaw, Bryan P. Glass, Rudolph J. Miller and William Warde have provided guidance, faith and great patience. Many other people have provid~d support and encouragement as well, particularly Drs. Tracy Carter and Helen Miller and many, many friends (especially Joe and Dixie), professors, colleagues and students along the way. For the loan of specimens or data I am most grateful to The Chicago Field Museum, The U.S. Museum of Natural History, The Museum of Natural History at the University of Kansas and the Stovall Museum at the University of Oklahoma. Finally, the ~reatest source of stren~th and consolation alon~ the way: to my ~arents, Keith and Betty, who tau~ht me determination. v TABLE OF CONTENTS Chapter Page I. INTRODUCTION 1 Justification • . • • • 3 Objectives .•. • 11 II. METHODS . • • 1 2 Site Descriptions • • • • • •••• 12 Regional..... • ••••••• 12 Vegetation ••••..••.•.••. 13 Climate • . • • • 15 Habitat Zones . • • • •••• 16 Field Techniques • . • • . 22 Small Mammal Identification •••••••. 28 Community Parameters • • • • • •••• 31 Number of Species (S) • • • • • • 31 Number of Individuals (N) • . • • 31 Index of Diversity (H') ••• 32 Evenness (J') .....•...... 32 Diversity (a) •••.•••••••• 33 Endemism (E) ••••.•••••••• 34 Dominance (D) • • • • •••. 34 Rarity Index (R) ••.•••••.•. 36 III. RESULTS . • • • • • • ••.•••••••• 37 Identification • • • • • • • • • • . 37 Small Mammals ••••••.•••.• 37 Large Mammals • • • • • • • • 64 Community Ecology • • •••••.••• 68 Small Mammals - Trapping Data •••• 68 Large Mammals - Transect Data •••• 71 Chiroptera - Mistnet Data •••. 76 Overall Community Parameters • • • • • 79 Seasonal Effects • , ••••••••••• 84 Large Ma~mals - Transect Data •••• 84 Small Mammals - Trapping Data •••• 90 Raptor Pellets • • • • • • • • • • 90 IV. DISCUSSION • • 93 Species Identification. • . ••• • 93 General Taxonomic Comments ••. • 93 Other Potential Caparao~Species • • • • 104 Altitudinal Affects on Community Ecology .108 vi Chapter Page Comparisons with Temperate Mountains •• • 111 v. SUMMARY . • • • • .118 VI. CONCLUSIONS AND RECOMMENDATIONS . .120 LITERATURE CITED . • 129 APPENDICIES . • • • • • . • .138 ,, APPENDIX A - MAMMAL SPECIES OF CAPARAO (IBDF) . .139 APPENDIX B - MAMMAL SPECIES OF CAPARAO " (RUSCHI) . .141 APPENDIX c - LIST OF SPECIMENS . .144 APPENDIX D - SPECIES LIFE HISTORY NOTES . .150 vii LIST OF TABLES Table Page 1. Descriptions and Effort for Data Collection Sites ••••••• . 23 2. Effort for Transects, Trapping and Mistnsts by Zone, Season, and Month • • • • • 25 3. P > F for General Linear Regression F-tests for Paired Species of Oryzomys • • • • • • • 40 4. Classification Matrix for Oryzomys Discriminant Analysis of 13 Oryzomys Species • • • • • • • • • • • • . • • • • 44 5. Southeastern Brazil Oryzomys Species Character Means and Ranges in Millimeters 49 6. P > F for General Linear Regression F-tests for Paired Species of Akodon • • • • • • 52 7. Classification Matrix for Discriminant Analysis of 12 Akodon Species •••• • 54 8. Southeastern Brazil Akodon and Related Genera's Species Character Means and Ranges in Millimeters • • • • • • • • • • • • 58 9. Cricetidae Other Than Akodon and Oryzomys Characters Means and Ranges in Millimeters • 59 10. Trapping Results of Small Mammal Species by Habitat (Excluding Recaptures) ••••••• 69 11. Mann-Whitney Results for Comparisons of Habitats Based on Small Mammal Species Ranked Frequencies From Trapping Data • ••••••• 72 12. Large Mammal Species Frequencies Per 100 Kilometers of Transect by Habitat •••••• 73 13. Mann-Whitney Results for Comparisons of Transect Data Between Habitat Zones •••• 77 viii Table Page 14. Bats Captured by Habitat Zone • • 78 15. Community Parameters for Caparao'Habitat Zone • 81 16. Dominance for Various Species Across Habitats. 83 17. Frequencies Per 100 Km of Transect for Nine Large Mammals by Habitat and Season • • 86 18. Chi - S4uare Values for Seasonal Changes in Large Mammal Frequencies (DF = 1) • • 88 19. Skulls Collected From Raptor Pellets by Season. 92 20. Comparisons of Community Parameters for Caµarao'and Great Smoky Mountains • .113 21. Linear Regression Correlations of Community Parameters with Altitudes for Caparao'and the Great Smokies • .114 22. Caµara~ Species Responses to Agriculture When Compared to Rainforest Based on Frequencies of Occurrence • .123 23. Summary of Numbers of Observations of Reproductive Data for Cricetidae .175 ix LIST OF FIGURES Figure Page 1. Map of Neotropical Refugia • • • • • • • • • • • • 5 2. Location and Map of Caparao National Park. Brazil. 14 3. Caparao National Park Weather Data at 1000 m. Elevation for 1980. • • • • • • • • • ••• 18 4. Altitudinal Variation in Ecological Zones of Caparao National Park • ••••••••••••• 19 5. Discriminant Function Plot of Eastern Brazil Cricetidae ••••••••••••••••••• 38 6. Discriminant Function Plot of 13 Oryzomys Species • • • • • • • • • • • • • • • • • • • • 45 7. Frequencies of Caparao Oryzomys Species According to Habitat ••••••••••••• 47 8. Discriminant Function Analysis of 5 Akodon Species • • • • • • • • • • • • • • • • 55 9. Akodon Species of Caparao by Habitat • . • • 5 7 10. Discriminant Function Plot of S.E. Brazil Genera Excluding Oryzomys and Akodon • • • • • • • • • 61 11. Monthly Glossophaga Population from Rock-slab Roost in Vale Verde • • • • • • • • • • • • • • 80 X' CHAPTER I INTRODUCTION The tropics contain the most diverse and, unfortunately, some of the most endangered terrestrial ecosystems on earth. Twenty-five to 50% of all the world's species occur in tropical forests (Ehrlich and Ehrlich 1981, Kaufman and Mallory 1989) and these forests are being lost at the rate of approximately 100 acres/minute, which had resulted in a 40% reduction from their original area by 1975 and an almost exponential rate of increase (or loss actually) projected over the next decade (Colinveux 1979, Kandell 1984). Recent satellite surveys indicate that the Amazon region had 178,000
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