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The Systematics, Biology, and Conservation of Solenodon

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THE SYSTEMATICS, BIOLOGY, AND CONSERVATION OF SOLENODON By JOSE ALBERTO OTTENWALDER A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1991 To Nicole To Alejandro To Ana Daniela ACKNOWLEDGEMENTS With justice, this could be another chapter. I thank my professors. Dr. John F. Eisenberg, Chairman, Dr. Charles A. Woods, Dr. F. Wayne King, Dr. Robert W. Woodruff, Dr. Mel Sunquist, and Dr. Brian McNab, for their guidance, support and advice. For their assistance, I am thankful to the people of the Parque Zoologico Nacional of Santo Domingo, the Florida Museum of Natural History, the Department of Wildlife and Range Sciences, the Instituto de Ecologia y Sistematica, Academia de Ciencias de Cuba, and the Museo Nacional de Historia Natural de Cuba. I thank the curators and staff of all of the natural history museums and zoological parks of North America, Europe and Caribbean that make specimens and information available to us. For their friendship and support, I thank Kent Vliet, Kevin Jordan, Laurie Wilkins, Gary Morgan, Dick Franz, and Julian Duval. I also thank Alfonso Ferreira, Angelica Espinal, and Roberto Maria in Santo Domingo, and Rosendo Martinez, Oscar Arredondo, Jose Fernandez Milera and Jorge de la Cruz in La Habana. I am grateful to Missy Woods, Perran Ross, and John Eisenberg for their hospitality in Gainesville. I thank Kent Vliet for his invaluable help with the graphics figures, and Lisa Chandler for the iii anatomical illustrations. For their wisdom and teachings, I thank the campesinos of the Dominican Republic. My studies and conservation work in the Caribbean was possible with grants from Wildlife Conservation International, division of New York Zoological Society, the Nixon Griffis Foundation, and Brehm Funds for International Bird Conservation. During my stay in Gainesville, I was supported with grants to Dr. John F. Eisenberg, Dr. Charles A. Woods, and the Program for Studies in Tropical Conservation. IV TABLE OF CONTENTS ACKOWLEDGEMENTS iii ABSTRACT vii CHAPTERS I INTRODUCTION 1 Evolutionary Relationships of West Indian Insectivores 3 Evolution of the Solenodontidae 6 Biology of Solenodon 9 II SYSTEMATICS OF THE GENUS SOLENODON 14 Materials and Methods 14 Results 24 Nongeographic Variation 24 Variation with age 24 Secondary sexual variation 25 Individual variation 26 Specific Relationships: Geographic Variation 27 Univariate analyses 27 Multivariate analyses 30 Variation in cranial morphology 33 Taxonomic Conclusions 39 Systematic Accounts 42 Genus Solenodon 42 Solenodon paradoxus 43 Distribution 43 Diagnosis 4 3 Comparisons 44 Geographic variation 45 Taxonomic conclusions 48 Solenodon paradoxus paradoxus 49 Solenodon paradoxus new subspecies B 52 Solenodon cubanus 57 Distribution 57 Diagnosis 57 Comparisons 58 Geographic variation 59 Taxonomic conclusions 59 v Solenodon new species A 65 Distribution 66 Diagnosis 66 Description 66 Comparisons 67 Solenodon marcanoi 67 Distribution 67 Revised diagnosis 67 Comparisons 68 Geographic variation 68 Taxonomic conclusions 70 III LATE QUATERNARY AND RECENT DISTRIBUTION 143 Material and Methods 143 Results 144 Solenodon paradoxus 144 Solenodon cubanus 147 Solenodon marcanoi 149 Solenodon "new species A" 149 IV BIOLOGY 160 Materials and Methods 160 Results 162 Reproduction and Development 162 Body Weights and Measurements 168 Thermoregulation and Diel Activity 170 Parasites 172 V CONSERVATION STATUS 195 Status and Numbers 195 Conservation Problems 200 Assessment of Captive Breeding 208 Conservation Measurements Taken 219 VI DISCUSSION AND CONCLUSIONS 236 LITERATURE CITED 261 APPENDICES 1 FOOD ITEMS AND DIET SUPPLEMENTS OF SOLENODON IN CAPTIVITY 277 2 PARASITISM AND PARASITE-RELATED MORTALITY IN SOLENODON PARADOXUS RECORDED IN POST-MORTEM EXAMINATIONS 278 3 SUMMARY OF HUSBANDRY PROCEDURES FOR CAPTIVE SPECIMEN OF S. PARADOXUS BETWEEN DECEMBER 1986 AND DECEMBER 1990 280 BIOGRAPHICAL SKETCH ori vi Abstract of Dissertation Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy SYSTEMATICS, BIOLOGY AND CONSERVATION OF SOLENODON By Jose Alberto Ottenwalder December 1991 Chairman: Dr. John F. Eisenberg Major Department: Forest Resources and Conservation An investigation of the geographic variation of Solenodon indicate that this Greater Antillean insectivore genus is represented by four species, two living, S. cubanus, S. paradoxus . and two extinct, S. marcanoi and a yet unnamed giant member from Cuba. The large fossil species from Cuba and a new geographic population of S. paradoxus from Hispaniola are described. The diagnosis of S. marcanoi is revised and some specimens of the original type series are re-assigned to S. paradoxus . S. marcanoi shares characters of both, S. paradoxus and S. cubanus . and is considered an intermediate lineage. Surveys indicate that S. paradoxus is dispersed widely in the Dominican Republic. The discovery of new populations of the species in that country is presented. Extant populations are fragmented in distribution and low in numbers. In Haiti, the species appear to survive only in vii . the southern Peninsula de Tiburon, yet it might exist in the Massif du Nord. Observations on the biology of S. paradoxus were recorded in the wild and in captivity. Males and females show reduced secondary sexual dimorphism in external morphology, and in cranial and post-cranial skeleton, though females are on average slightly larger and heavier. The permanent dentition emerges completely by 4-5 months of age. Males and females attain adult size before eight months and might reach sexual maturity soon after one year of age. Age at first reproduction is probably around 18 months. Gestation lasts for about three months, and litter size is one. Body temperature is low, ranging from an average 33 °C during daytime to a peak of 36 °C during nocturnal foraging. Limitations in thermoregulatory ability might be compensated by the microclimatic stability of the burrow. Diel activity patterns show a sleeping-resting phase throughout the day, which ends with a sudden start of activity at or after 1800 h. Nocturnal activity is characterized by successive foraging trips at variable intervals, with peaks between 2000-2400 h. Crepuscular activity is less freguent. The residual populations of S. cubanus in eastern Cuba appear to be critical. Systematic efforts and direct conservation action would be required to ensure their survival viii . CHAPTER I INTRODUCTION The West Indian insectivores, Solenodon and Nesophontes . are endemic to the Greater Antilles and the most ancient members of the West Indian mammalian fauna. The genus Solenodon is restricted to the islands of Cuba and Hispaniola and contains the only surviving members of the Insectivora in the region. The closely related Nesophontes comprises eight extinct species known from the Holocene and Late Pleistocene of Cuba, Hispaniola, the Cayman Islands and Puerto Rico (Fig. 1-1) . The Puerto Rican species, N. edithae . which is intermediate in size between the larger Solenodon and the much smaller remaining species of Nesophontes . has been recently discovered in a kitchen midden in Vieques Island (Morgan and Woods 1986, E. Wing pers . comm. ) Though they have fared better than other groups such as edentates and primates, insectivores have also suffered the high extinction rates recorded among other West Indian mammals (Morgan and Woods 1986, Woods 1989, 1990). Two of twelve species of insectivores have survived until today. Pleistocene climatic events, human exploitation and predation pressure from exotics have been indicated as major 1 2 causes of extinction of the Antillean vertebrate fauna. Increasing support for the latter two factors have been presented (Steadman et al. 1984, Woods et al. 1986, Woods 1989) . Most West Indian mammals were still extant at the time Amerindians arrived on the islands (Morgan and Woods 1986) . In fact, all three species of Hispaniolan Nesophontes might have survived to the beginning of this century (Miller 1929) . Both Nesophontes and Solenodon have been found in archaeological sites throughout their historical range. Their presence in archaeological deposits is, however, virtually insignificant compared to the abundance of other groups, and they do not seem to have represented an important source of human food. Evidence from cave deposits and owl pellet accumulations indicate that Nesophontes were clearly very abundant, but no Solenodon . Therefore, Nesophontes were undoubfully neglected as food by the Amerindians because of their small body size. Although Solenodon species were much larger, they do not seem to be more common in archaeological sites, and they are infrequent in cave deposits. Predation from exotics appears to be the single most important cause of their extirpation. Today, Cuban and Hispaniolan Solenodon are among the few native West Indian land mammals that still survive. They are considered among the most endangered mammals, and probably are the most threatened of all insectivores (Thornback 1983; Thornback and Jenkins 1982). 3 Evolutionary Relationships of West Indian Insectivores The early evolutionary history of the group, was comprehensively summarized by McDowell (1958) . The relationships of Solenodon and Nesophontes . among themselves and within the Insectivora, are not yet well understood. In part, this is a reflection of the continuing problems
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