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I J?) 8 (i|' #] I • ‘ 2 to ‘ 2 to 2 ^ ^ HoiiniiiSNi_NviNOSHims saiyvy SNi ~ NviNOSHiiiMS S3iyvyan libraries Smithsonian institution to 2 to — 2 \ ^ ^ tiJ UJ UJ 2 c/, V to tj . r? o o “ LIBRARIES SMITHSONIAN INSTITUT ES SMITHSONIAN INSTITUTION NOIiniIiSNl”'NVINOSHiIlAIS S3 I 8 V 8 9 IT \<2jcdcJ to ± ^ to \ ^ CO ± to LSNI NoiiniiisNi NviNOSHims NviNOSHims ssiyvyaiT libraries Smithsonian institution ~ "* S3iyvy to CO 2 2 ..•• 2 ,,, to _ -. ^-2 W *. 2 to 2 I LI B RAR I ES^SMITHSONIAN_ INSTITUT ES^^SMITHS0NIAN_ INSTITUTION NOIiniIlSNI_NVINOSHiIlNS S3 I 9 Vy g n_ <" A,«wv.\ - /iSSx S i ^ - 111) r! ^2^ 5^^ a , //£ c: ~ xc O — '<ioxu2^ Q 'X '^^ojuAjX O y< pc^ " O LSNl"^NVINOSHimS S3 I H VU 9 IT "*LI B R A R I ES^SMITHS0NIAN“'|NSTITUTI0N NOliniliSNl'^NVINOSHiU^S S3iyvy f- , 2 r~ 2 2 r”, 2 o — ><5'al^x O m 2 m ^ 2 BERYL B. SIMPSON 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 Sm/thson/an 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 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. S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 43 A Revision of the Genus Polylepis (Rosaceae; Sanguisorbeae) Beryl B. Simpson ISSUED .NOV.271979 SMITHSONIAN PUBLICATIONS SMITHSONIAN INSTITUTION PRESS City of Washington 1979 — ABSTRACT Simpson, Beryl B. Revision of the Genus Polylepis (Rosaceae: Sanguisorbeae). Smithsonian Contributions to Botany, number 43, 62 pages, 39 figures, 1979. Species of Polylepis are shrubs or trees native to the mid- and high-elevation tropical Andes. Some species of Polylepis form woodlands growing well above normal treeline within grass and scrub associations at elevations over 5000 m. Consequently, Polylepis appears to be the highest natural occurring arborescent angiosperm genus in the world. The physiological basis allowing such high alti- tude growth is not understood, but evidence indicates that woodlands at these elevations are restricted to microsites where ecological conditions are similar to those of lower altitudes. The genus is distinct from other rosaceous members of the tribe Sanguisorbeae and is interpreted here as consisting of three species groups that appear to have spread independently north and south along the Andean chain. Because of the patchy distributions of populations and disturbances by man, species are often polytopic. As a result, numerous forms have been de- scribed as distinct taxa in the past. In addition, hybridization appears to occur between species making circumscription of some taxa difficult. In this revision, evidence from studies of gross morphology, leaflet anatomy, pollen morphology, and field observations of the autecology of various populations were used to ascer- tain the number of species within the genus and their relationships to one another. Data from these sources indicate that Polylepis should be considered as consisting of 15 species, one of which is first described in this treatment. A key for the iden- tification of the species is provided. Each species is illustrated, discussed, and its modern distribution mapped. Official publications date is hanclstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. Series cover design: Leaf clearing from the katsura tree Cercidiphyllnm japonicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data Simpson, Beryl B. A revision of the genus Polylepis (Rosaceae: Sanguisorbeae) (Smithsonian contriliutions to botany ; no. 43) Bibliography: p. 1. Polylepis. 2. Botany-—-Andes. 1. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to botany 43. ; no. QK1.S2747 no. 43 [QK495.R781 581'.08s [583'.372] 79-986 Contents Page Introduction I Acknowledgments Previous Treatments of the Genus Species Groups and Relationships of the Genus 3 Morphological Characters 4 Habit 4 Bark 4 Branching Pattern and Leaf Arrangement 6 Stipule Sheaths 6 Trichome and Vestiture Types h Leaves and Leaflets . 7 Leaf Anatomy 8 Inflorescences and Flowers 12 Pollen Morphology 14 Fruits 14 Geographical Distribution 14 Ecology 15 Polylepis Ruiz and Pavon 17 Key to the Species of Polylepis 18 1. Polylepis multijuga Pilger 19 2. Polylepis lanuginosa Humboldt, Bonpland, and Kunth 21 3. Polylepis hieronymi Pilger 23 4. Polylepis pauta Hieronymus 27 5. Polylepis sericea Weddell 28 6. Polylepis subsericans Macbride 31 7. Polylepis pepei, new species 32 8. Polylepis reticulata Hieronymus 33 9. Polylepis weberhaueri Pilger 37 10. Polylepis quadrijuga Bitter 40 11. Polylepis besseri Hieronymus 42 12. Polylepis tornentella Weddell 46 13. Polylepis incana Humboldt, Bonpland and Kunth 50 14. Polylepis racemosa Ruiz and Pavon 53 15. Polylepis australis Bitter 56 Literature Cited 60 Index 62 iii r :, ^;b ^ - ‘ ED J . -' ^.Ovv.(,'N>ln,! . , , - - » 'J*.^ 4A k I r *- ( Ifl UMk ^ , . I .k# v)i j>rr» iV '« ' i I < '», _ ^ .%T^ Jr t.'- ‘ . ' jk' t Jtj'.jj '> , I' ',1 !i ht!^ iO ..^:'!b •• |ti Jti t ^ |rt) Cfi I . il I I .'• kuK i... “'> -I... /I & ’ * iitl ' ' ^/-•J,". ' -i M -Ik' '*>7 "- *' 4!^^^^]! kUfj)/ :^,v:. 'iU ',•'••'• '.'« »'7 ’ t ’» _jM‘ '.K * I I »yt•^,l^^lJ. li' 'k'.,',V,J .;t'j*l<('Wt ' I ' ‘ ' •< **'{<<%* ' •..n- I i.M 'J ’. , v''"5k 'k'l Si... i m .v-tiiir-. ^S' ..V.' ii : A\ A Revision of the Genus Polylepis (Rosaceae: Sanguisorbeae) Beryl B. Simpson Introduction Above elevations of 3500 m in the tropical Andes, erable debate, but evidence indicates that even be- vegetation usually consists of various forms of low fore severe decimation by man, high elevation trees grass and scrub paramo or puna. Woody elements of Polylepis were limited in their distribtution by are scarce at such elevations in terms of both num- the presence of specialized microhabitats. bers of species and individuals. Yet, one arborescent In addition to its uniqueness as an inhabitant of genus, Polylepis, is found throughout the high extremely high elevations, Polylepis has played an tropical Andes.