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Swollen-Thorn Acacias of Central America

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Swollen-Thorn Acacias of Central America SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 13 Swollen-Thorn Acacias of Central America Daniel H. Janzen SMITHSONIAN INSTITUTION PRESS City of Washington 1974 ABSTRACT Janzen, Daniel H. Swollen-Thorn Acacias of Central America. Smithsonian Contributions to Botany, number 13, 131 pages, 119 figures, 1974.-This nomen- clatural, taxonomic, and ecological treatment of 11 Central American obligate ant-acacias (Acacia allenii, A. chiapensis, A. collinsii, A. cookii, A. cornigera, A. gentlei, A. globulifera, A hindsii, A nzayana, A. melanoceras, and A. sphaero- cephala) and one quasi-obligate ant-acacia (Acacia ruddiae) is based on ex- tensive field study from 1963 to 1972 and on herbarium specimens where of use. 7 he population boundaries of all species are mapped and described with respect to ecological parameters. Morphological variation, details of the interaction with the ants, and acacia reproductive biology are presented for most species. OFFICIALPUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution‘s annual report, Srnithsonian Year. SI PRESSNUMBER 4817. SERIESCOVER DESIGN: Leaf clearing from the Katsura tree Cercidiphyllurn japonicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data Janzen, Daniel H: Swollen-thorn acacias of Central America. (Smithsonian contributions to botany, no. 13) Bibliography: p. 1. Acacia. 2. Ants-Central America. 3. Botany-Central America-Ecology. I. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to botany, no. 13. QKl.S27 no. 13. [QK495.L52] 581’.08s [583’.321] 73-4401 For sale by the Superintendent of Documents, US. Government Printing O5ce Washington, D.C. 20402 . Price $2.35 (paper cover) Contents Page Introduction ................................................... 1 General Biology ................................................. 2 General Comments on Swollen-Thorn Acacia Taxonomy .............. 6 Materials, Methods, and Terminology ............................... 14 General Morphological Traits .................................. 15 Taxonomic Information .............. ....................... 25 A . Key to Mature Central American Swollen-Thorn Acacias, Based on All Traits ........................................... 28 B . Key to Mature Central American Swollen-Thorn Acacias, Based on Vegetative Traits .................................. 29 Acacia cookii Safford ................. ................... 29 Acacia ruddiae, new species ..................................... 34 Acacia melanoceras Buerling ..................................... 42 Acacia allenii, new species .......................................... 53 Acacia chiapensis Safford ....................................... 61 Acacia globulifera Safford ........................................ 67 Acacia sphaerocephala Schlectendal and Chamisso ..................... 73 Acacia cornigera (L.)Willdenow .................................. 80 Acacia mayana Lundell ........................................... 93 Acacia gentlei Standley ....................................... 95 Acacia collinsii Safford ......................................... 99 Acacia hindsii Bentham ........................................ 113 Literature Cited ................................................. 130 Swollen-Thorn Acacias of Central America Daniel H. Janzen Introduction len-thorn acacias, a stable nomenclature is essential at this time. (3) Detailed data on populations of na- The swollen-thorn acacia group (Acacia species) tive wild Central American plants are noticeably is one of the two most diverse and widespread of absent from the literature, A better concept is the neotropical myrmecophytes. These acacias are needed of the population structures of a wide va- the characteristic ant-plants in the drier lowlands riety of tropical plants of different life forms and in (“tierra caliente”) of Central America, just as the different habitats in order to generate specific hy- genus Cecropia (Aloraceae) is the most conspicuous potheses that can lead to an understanding of com- and diverse myrmecophyte of wetter habitats in plex tropical communities. (4) The lowland Central and South America. deciduous forests of Central America have nearly This monograph on the taxonomy and natural disappeared. The population structures of the spe- history of neotropical swollen-thorn acacias is of- cies remaining in these habitats are already greatly fered at this time for several reasons. (1) The altered. The swollen-thorn acacias are clearly in this neotropical swollen-thorn acacias do not appear to category. I would like to summarize the ecological constitute a tight phjletic unit, yet they share many behavior of swollen-thorn acacias before their pat- adaptive ecological and morphological traits. They terns are so far altered that we can no longer recog are thus outstanding examples of evolutionary con- nize the conditions that produced the genotypes we vergence. The most noteworthy of the traits they find in today’s cattle pastures, roadsides, and have in common is that they are protected from abandoned fields. Most wild plants in high-yield herbivores by colonies of ants in the genus Pseudo- agricultural areas must be studied under conditions myrmex, rather than by chemical traits as are most not unlike studying elephants and deer in a zoo. plants (Janzen, 1966a, 1967b; Rehr et al., 1973). (2) It is hoped that the incompleteness of the data The ants can be readily removed, just as though and hypotheses offered here will encourage field one were able to easily remove the alkaloids and biologists to look harder at why plants do what they cyanogenic glycosides from the leaves of ordinary do in natural habitats as well as in those badly acacias such as Acacia farnesiana. This ease of ex- altered by agriculture. perimentally disarming the plant means that the ACKNOWLEDGMENTS.-~am greatly indebted to protective role of the obligate acacia-ants has been Dr. Velva E. Rudd of the National Museum of easy to study and provides an excellent system for Natural History for her very competent assistance understanding the coevolution of complex interac- and advice throughout this study. Her patience tions between higher organisms. For further and with the bumblings of an entomologist in the world more detailed comparative studies among the swol- of taxonomic botany immensely facilitated the un- Daniel H. Janzen, Department of Zoology, University of Mich- raveling of the nomenclatorial spaghetti surround- igan, Ann Arbor, Michigan 48105. ing the swollen-thorn acacias. Personnel in various 1 SMITHSONIAX CONTRIBUTIOXS TO BOTANY herbaria have kindly allowed me to examine types Acacia are primarily in areas with less than 3 me- and at times to borrow specimens. ters of annual rainfall and with a well-developed A special acknowledgment is due Karen Janzen, dry season of 3-11 months duration. Throughout who participated in the laborious task of recording Africa and the neotropics, those species with foliar field data and voucher specimens. nectaries (probably all species) have a loose associ- This study was supported by National Science ation with ants. Many genera of arboricolous ants Foundation Grant GB-5206, and is in part a by- (e.g., Crematogaster, Pseudomyrmex, Camponotus) product of NSF Grants GB-91, 1428, 7805, and visit these nectaries and in the process probably in- 7819. Substantial portions of this study were com- crease the intensity of their foraging activity on pleted while I was serving in Costa Rica as a faculty these plants compared with adjacent plants lacking member for the Organization for Tropical Studies nectaries. This, accompanied by similar benefits in Costa Rica (1965-1971). The paper is Contribu- from parasitic Hymenoptera attracted to the nec- tion No. 1360 from the Department of Entomology, taries, has probably been the selective force leading The University of Kansas, Lawrence, Kansas. to the evolution of these nectaries (cf. Janzen, 1966a; Bequaert, 1922). In a few cases, however, the ant and acacia inter- General Biology action has evolved to a much higher level of com- The 12 species of swollen-thorn acacias in aggre- plexity, that of obligatory mutualism (Janzen, gate range from the Tropic of Cancer on both 1966a). For background information, a brief resume coasts of Mexico only as far south as northwestern of the interaction in Central America is given in Colombia. They are generally found at elevations the following paragraphs. More details can be ob- below 1200 m (Figure 1). One Argentinian acacia, tained from Janzen’s (1967b) lengthy description of Acacia cauen (Molina) Molina, has been suggested Acacia cornigera and Pseudomyrmex ferruginea in as a swollen-thorn acacia (Fiebrig, 1909; Chodat lowland eastern Mexico and in peripheral papers and Vischer, 1920); however, Wheeler (1 942), on ants and acacias (Janzen, 1967a, 1969a, 1969b, Havrylenko (in litt.), and Sliillink (in litt.) have 1973; Beulig and Janzen, 1969). convinced me that the ants living in the thorns A colony of obligate acacia-ants lives in the swol- were only fortuitous invaders. My own unpublished len, stipular thorns of one to several individual observations of A. cazlen in Chile give no hint of shrubs or small trees that have been given the sub- myrmecophytism with this plant. Greenhouse A. generic common name of “swollen-thorn acacia” cazlen (from Argentinian seeds) shows thorn and (Janzen, 1967a). New World obligate acacia-ants leaf development like that of normal acacias and are a behaviorally
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