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A Taxonomic Study of the Genus Eugenia (Myrtaceae) in Hawaii1

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A Taxonomic Study of the Genus Eugenia (Myrtaceae) in Hawaii1 A Taxonomic Study of the Genus Eugenia (Myrtaceae) in Hawaii1 KENNETH A. WILSON2 THE PANTROPIC GENUS Eugenia (Myrtaceae) taken this problem under his guidance and is represented in the forests of Hawaii by am indebted to him for his help and en­ several species. A recent taxonomic treatment couragement. I wish to thank Miss Marie C. of the genus by Merrill and Perry (1939) Neal, Curator of the Herbarium of the Bernice divided the greater number of species of P. Bishop Museum, for her helpful and valu­ Eugenia into the genera Syzygium and Eugenia able assistance and for the opportunity to sensu strictu. A later study by Henderson work on the specimens in the herbarium. Dr. (1947), based on the Malayan species, re­ Rogers McVaugh, Curator of Phanerogams jected this classification. of the Herbarium of the University of Michi­ According to the Merrill and Perry classi­ gan, has generously made the space and fication, both Syzygium and Eugenia sensu facilities of the herbarium available to me for strictu are present in the Hawaiian flora. There­ continuing this study. I gratefully acknowl­ fore, it seemed that an evaluation of the re­ edge the assistance of Dr. Joseph F. Rock, cent generic treatments based on these who provided valuable information based on representatives would be of value. My study his knowledge and experience with the Ha­ of the generic status of Eugenia and Syzygium waiian flora. led to the question of the specific status of the I am indebted to the following herbaria for indigenous Hawaiian members. As a result the loan of herbarium specimens, photo­ of my investigations I have rejected the trans­ graphs of types, and other valuable materials: fer of species of Eugenia to Syzygium, de­ Herbarium of Bernice P. Bishop Museum, scribed a new species from the island ofMolo­ Gray Herbarium, Herbarium of the Royal kai, and reduced two species and one variety Botanical Gardens at Kew, Botanische Staats­ to synonomy. sammlung Mlinchen, Herbarium of the Uni­ versity of Michigan, Herbarium of the New ACKNOWLEDGMENTS York Botanical Gardens, and United States National Herbarium. Prof. Harold St. John suggested to me that the problem of the generic status of Eugenia HISTORY OF THE GENUS was in need of further study. I have under- Since Linnaeus' treatment of Eugenia in Species Plantarum (1753: 470), more than 800 1 Part of a thesis submitted to the Graduate School of the University of Hawaii in partial fulfillment· of the species have been described or transferred to requirements for the degree of Master of Science. this genus. Many botanists have been dis­ 2 Department of Botany, University of Michigan, Ann Arbor, Michigan. Manuscript received February satisfied with the wide range of difference in 10, 1956. form shown by the members of the genus. 161 162 PACIFIC SCIENCE, Vol. XI, April, 1957 One of the most comprehensive reclassifica­ intergrading series from those with two dis­ tions of Eugenia was proposed by Niedenzu tinct and separate cotyledons to those with (1893). He established several segregate gen­ the cotyledons completely fused. On the basis era which were based mainly on the char­ of these observations he rejects the transfer acters of the Rower. His system was adopted of species from Eugenia to Syzygium. by many workers, but, because of the lack of In 1953 Ingle and Dadswell published the distinct generic limits, many other botanists results of their studies of the wood anatomy continued to consider Eugenia in a broader of several Pacific M yrtaceae. 3 They concluded sense. Later studies of tbe members of the that the anatomy of the wood of Eugenia group led more and more botanists to reject provides ample characters for splitting the Niedenzu's classification and to return the genus. They added that the "suggested split segregated genera to Eugenia. on anatomical grounds involved onlY,the two Because of the great number of species groups Eugenia A [corresponding to Eugenia described in Eugenia the group has become sensu strictu] and Eugenia B [including the rather unwieldy, and Merrill and Perry (1938a, genera Acmena, Cleistocalyx and Syzygium]."4 1938b, 1939) proposed a new systematic The most recent treatments of the genus treatment of the group. They redefined some have been inconsistent. By far the majority of of the earlier proposed segregates of the workers (Airy Shaw, 1949; Backer, 1945; genus. The new systematic treatment which Degener and Ludwig, 1952) have accepted they proposed is based on the structure of the Merrill and Perry's classification without any seed. According to them, species of Eugenia apparent critical study. A few botanists are diagnosed as having a pericarp which is (White, 1945; Amshoff, 1942), seemingly in easily crushed and "the seed is free, the testa doubt as to the validity of this system, have is smooth, chartaceous to cartilaginous and hesitated to adopt it. mostly lustrous, and the cotyledons are me­ chanically inseparable, i.e., they have grown EVALUATION OF THE RECENT together in such a way that often the line of GENERIC TREATMENTS their opposing faces is scarcely distinguish­ The Hawaiian Rora, although predomin­ able." Syzygium, one of the segregate genera, antly Asiatic in its affinities, includes repre­ is described as having "fruits that when dried sentatives of the tropical regions of both the are not too easily broken, and, when opened, Old World and the New World. Because of the embryo (not the entire seed) falls out the uniqueness of the Island Rora in this leaving the roughish seed coat more or less respect, the Hawaiian Islands may be con­ loosely adhering to the pericarp; the embryo sidered an ideal region for evaluating the re­ has two distinct cotyledons usually attached cent treatment of Eugenia. near the middle of the opposing faces which The two most important features used by conceal the hypocotyl within." Merrill and Perry in segregating the genera In a monograph of the species of Eugenia are: (1) the nature of the embryo, and (2) the in Malaya, R. M. Henderson (1949) critically degree of adherence of the testa to the coty­ analyzed Merrill and Perry's classification. ledons. On the basis of these characters Henderson, in his study, decided that neither the degree of adherence of the seed coat to 3 See also Dadswell and Ingle, 1947. 4 Since this paper went to press Kathleen M .. Pike the cotyledon, nor the nature of the cotyle­ published the results of her studies in the pollen dons themselves is consistent enough to be morphology of the Myrtaceae. (Austral. Jour. Bot. regarded as a good generic character. In fact, 4(1): 13-53, 1 pI., 1956.) She found that the pollen grains of "Eugenia A" are distinct from those of he gives detailed descriptions of the seeds of "Eugenia B" and thereby adds support to the group­ several different species which illustrate an ings of Ingle and Dadswell based on the wood anatomy. Eugenia in Hawaii - WILSON 163 Merrill and Perry (1938a, 1938b, 1939), in dried and fresh fruits the thin seed coat peels studies of Indo-Chinese, Chinese, and Bor­ off with the pericarp. nean species of Eugenia, redefined the group, On the basis of the studies I have made, it placing most of the species of those regions seems evident that neither the character of the in the genus Syzygium. Geographically, their seed coat nor that of the embryo offers a new concept limits Eugenia mainly to tropical satisfactory basis for the reclassification of the America, and Syzygium primarily to the tropics group. Neither one of the characters is con­ of the Old Wodd. stant. The degree of fusion of the cotyledons The genus Eugenia in the Hawaiian Islands varies even within a single species. is represented by eight species as I interpret The conclusions I have drawn on the basis them. In my opinion, according to the Merrill of investigations of the Hawaiian representa­ and Perry system, four of the Hawaiian rep­ tives of Eugenia strongly support Henderson's resentatives of Eugenia would be classified in rejection of the segregation of species of the segregate genus Syzygium, while four Eugenia into the genera Syzygium and Eugenia would fall in Eugenia sensu strictu. sensu strictu. Eugenia malaccensis and E. Jambos have No attempt was made to study the wood fruits which contain a rather large seed loosely anatomy of the Hawaiian species of Eugenia. attached within the pericarp. In both species In view of Ingle and Dadswell's findings the embryo has two distinct, fleshy cotyle­ (1953) I suggest that additional more exten­ dons. The surface of the cotyledons is very sive investigations may reveal other morpho­ rugose and the thick seed coat adheres firmly logical characters to support their conclusions. to it. Young seedlings of E. malaccensis were The characters of the seed, since they are so found growing which had the two cotyledons variable, do not justify the split. still attached. The pericarp had rotted a";ay and no evidence of it could be seen, but on VARIA nON AND DISTRIBUTION OF THE SPECIES the surfaces of the cotyledons remains of the IN HAWAII seed coat were still present. In fruits of Eugenia Cumini the fleshy peri­ The genus is represented in Hawaii by only carp peels off, leaving the seed coat firmly four indigenous species, three of which are adhering to the cotyledons. The cotyledons endemic. Of these four, Eugenia rariflora and are fleshy and distinct. In dried, raw fruits the E. sandwicensis .show the greatest degree of seed coat readily peels off with the pericarp. variation within the entire group. Both show However, when boiled, the pericarp may be variations in leaf size ranging from very small easily removed without disturbing the seed to rather large.
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