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Holocene Vertebrate Fossils from Isla Floreana, Galapagos

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Holocene Vertebrate Fossils from Isla Floreana, Galapagos Holocene Vertebrate Fossils from Isla Floreana, Galapagos DAVID W. STEADMAN SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 413 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 413 Holocene Vertebrate Fossils from Isla Floreana, Galapagos David W. Steadman SMITHSONIAN INSTITUTION PRESS City of Washington 1986 ABSTRACT Steadman, David W. Holocene Vertebrate Fossils from Isla Floreana, Galapagos. Smith- sonian Contributions to Zoology, number 413, 103 pages, 25 figures, 4 plates, 12 tables, 1986.—This study surveys the late Holocene vertebrate fossil record from Isla Floreana, Galapagos. Over 20,000 fossils from four lava tubes in the arid lowlands near Post Office Bay are associated with six radiocarbon dates of 2400 years B.P. or younger. The fossils, most of which originated as regurgitated pellets of barn owls, represent more than 1100 individual animals of 24 indigenous species. They include six species now extinct on Floreana: Geochelone elephantopus (tortoise), Alsophis biserialis (snake), Tyto punctatissima (Galapagos Barn Owl), Mimus trifasciatus (Floreana Mockingbird), Geospiza nebulosa (Sharp-beaked Ground Finch), and Geospiza magnirostris (Large Ground Finch). These species are, respectively, 1 st, 7th, 16th, 6th, 15th, and 2nd in abundance among all species recorded as fossils, making up 57% of the individuals in the fossil fauna. In addition, the 3rd and 4th most common fossil taxa, Tropidurus grayii (lava lizard) and Zenaida galapa- oensis (Galapagos Dove), are extremely rare today on Floreana. Thus, extinction probably f ias changed the composition of Floreana's fauna even more than is suggested by the number of extinct species alone. While the e\idence is circumstantial, I believe that all extinction on Floreana is related to human impact, directly through predation and habitat alteration, and indirectly through the effects or alien animals (rats, mice, cats, dogs, pigs, goats, cattle, and donkeys). People and feral mammals have lived on Floreana since 1832, and most or all vertebrate extinction occurred within the succeeding 40-50 years. Direct predation by man and introduced mammals was probably the main cause of extinction only for Geochelone elephantopus, although such predation may have been involved to some extent in each of the other extinctions. Loss of preferred prey species and human predation probably caused the extinction of Tyto punctatissima. Extinction of Mimus trifasciatus and Geospiza magnirostris may have resuftea from destruction of Opuntia cactus by feral herbivores. Extinction of Geospiza nebulosa may be related to habitat changes in the highlands. Buteo galapagoensis (Galapagos Hawk), though not recorded as a fossil, is also apparently extinct on Floreana, with direct human predation likely to be the main cause. All extinction on Floreana occurred in historic times; whether this is true elsewhere in the Galapagos awaits more research. Lasiurus borealis (Red Bat) is the only indigenous mammal recorded as a fossil. The absence of fossils of cricetine rodents or Conolophus spp. (Land Iguana) is evidence that these forms never occurred on Floreana. The lack of fossils of Coccyzus melacoryphus (Dark-billed Cuckoo) and Dendroica petechia (Yellow Warbler), which occur commonly on Floreana today, is evidence that these two species colonized the Galapagos very recently. Fossil records depend upon suitable environments of deposition. The Galapagos Islands, being of recent volcanic origin, lack many potentially fossiliferous geological features such as fine-grained alluvial sediments, indurated sand dunes, or limestone caves and sinkholes. Fortunately, lava tubes, which are very well suited for accumulation and preservation of vertebrate fossils, are common on certain islands in the Galapagos. Fossils enable us to reconstruct undisturbed (pre-human) insular faunas more com- pletely than previously possible. Modern biogeographical studies usually do not consider now natural the faunas are; they would benefit by considering changes wrought by human impact. The field of island biogeography woula profit from a renewed emphasis on the collection of original field data rather than the current trend of theoretical studies that often are based upon inadequate or poorly understood data. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging in Publication Data Steadman, David W. Holocene vertebrate fossils from Isla Floreana, Galapagos. (Smithsonian contributions to zoology ; no. 413) Bibliography: p. Supt. of Docs, no.: SI 1.27:413 1. Vertebrates, Fossil. 2. Paleontology—Recent. 3. Paleontology—Galapagos Islands— Santa Maria Island. I. Title. II. Series. QL1.S54 no. 413 [QE881] 591 s [566] 85-600002 Contents Page Introduction 1 Abbreviations 1 Acknowledgments 2 Materials and Methods 3 Field Work 3 Museum Research 4 Modern Specimens Examined 5 Place Names 5 Background Studies 6 Geology 6 Climate 10 Paleoclimate 12 Vegetation 13 The Fossil Sites 15 The Caves 15 Stratigraphy 20 Chronology 25 The Fossil Fauna 27 Systematic Paleontology 37 Discussion 60 Human History 60 Introduced Mammals 61 Extinction 63 Theoretical Biogeography and Fossils 76 Animals Not Recorded As Fossils 83 The Paleontological Potential of the Galapagos 85 Summary and Conclusions 86 Literature Cited 89 Plates 100 in Holocene Vertebrate Fossils from Isla Floreana, Galapagos. David W. Steadman Introduction studies of the fossils from the other four islands Biologists have long held the Galapagos Islands are still incomplete. The fossils described herein, in high esteem. Even non-biologists are im- none of which predates with certainty even the pressed by the living things they see there, espe- Holocene, provide critical new evidence for un- cially by the tameness of the native animals. derstanding the extinction and biogeography of Situated in the equatorial Pacific Ocean, the the vertebrates of Floreana and the entire Gala- Galapagos Islands possess a terrestrial flora and pagos Archipelago. fauna that is derived almost entirely from the ABBREVIATIONS.—Terminology used through- South American mainland, nearly 1000 km to out this work includes the following: the east. More than any other locality in the B.P. before present (= before A.D. 1950) world, these inhospitable volcanic islands have CDRS Charles Darwin Research Station, Santa Cruz, Galapagos been regarded as a Mecca to which many natural ITCZ Intertropical convergence zone historians sooner or later pay homage. Evolution- MNI minimum number of individuals ary biologists, biogeographers, and ecologists have found the Galapagos particularly enticing The following persons, listed in Table 1, as- as an unparalleled "natural laboratory." In at- sisted in field work on Floreana: tempting to reconstruct the evolution of the ver- MJC Maria Jose Campos tebrates that inhabit the Galapagos, biologists GD Gayle Davis DG David Graham have lacked any paleontological evidence. Dur- HH Harvey Helman ing five trips to the Galapagos since 1978,1 have JRH James R. Hill, III collected fossils of reptiles, birds, and mammals GM Godfrey Merlen from lava tubes on five of the larger islands in MP Miguel Pozo the group—Santa Cruz, San Cristobal, Floreana, DWS David W. Steadman Rabida and Isabela. These specimens represent ENS Edward N. Steadman the first serious paleontological effort to docu- For species accounts in the "Systemic Paleon- ment the evolution, past distribution, and extinc- tology" section, the following acronyms are used tion of vertebrates
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