A Taxonomic Study of the () in Hawaii1

KENNETH A. WILSON2

THE PANTROPIC GENUS Eugenia (Myrtaceae) taken this problem under his guidance and is represented in the forests of by am indebted to him for his help and en­ several . 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 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 , 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 , 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 . 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 " that when dried sentatives of the tropical regions of both the are not too easily broken, and, when opened, and the . 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 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 ­ 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 size ranging from very small easily removed without disturbing the seed to rather large. coat. The greatest degree of variation is seen in Upon removal of the pericarp the testa of Eugenia sandwicensis. The forms of E. sand­ Eugenia sandwicensis also remains attached to wicensis with the larger are found most the cotyledons. The two fleshy cotyledons often on Kauai; the forms with elliptical and are not consolidated. In boiled dried material, elliptical-lanceolate leaves are more prevalent depending on the amount of care given it, the on Maui, and most of the Oahu representa­ pericarp mayor may not peel away from the tives have the smaller obovate leaves. On seed coat. Lanai the majority of the representatives of The seeds of Eugenia rariflora and E. koolau­ this species have small obovate leaves similar ensis have their cotyledons plainly separated, to those of the most common forms on Oahu. partly fused, or entirely consolidated. When Most of the members on have leaves the cotyledons are consolidated the line of which resemble the forms on Maui, although their opposing faces is not evident. In both in general they are not as long. It is possible 164 PACIFIC SCIENCE, Vol. XI, April, 1957

for a person who is familiar with the species to determine, in the majority of cases, from Eugenia L., Sp. PI. 470, 1753. which one of the islands a specimen was collected. However, this broad generalization Syzygium Gaertn., Fruct. 1: 166, t. 33, 1788. of geographical distribution is not constant, Jambosa DC., Prodr. 3: 286, 1828. and almost all forms are present on each of the or . Shoots glabrous or pube­ islands where the species occurs. scent. Leaves simple, opposite, glandular­ Of the naturalized species Eugenia Cumini punctate, pinnately veined with a continuous is by far the most widely distributed. E. intramarginal vein. Leaf scar with a single malaccensis is confined mostly to shaded, vascular bundle. single or in pairs, moist valleys where it is likely they were axillary, or in terminal, or axillary cymes or planted originally by the Hawaiians. Both racemes or inflorescence on leafless branches. Eugenia Jambos and E. uniflora are only sparsely Calyx t~be5 globose to elongate-turbinate, distributed in the native forests. extending beyond the ovary or not so, usually These introduced species are remarkably minutely glandular-punctate; calyx lobes 4, constant in their characters and show no sig­ large persistent and spreading, or small and nificant degree of variation. early deciduous; petals 4, free and spreading EXPLANA nONS persistent or caducous, or cohering and falling off as a calyptra; numerous, free on a Unless otherwise indicated, the specimens staminal disk lining the calyx tube or inserted studied are in the Bernice P. Bishop Museum. on the margin of the calyx tube; anthers versa­ The standard abbreviations of Lanjouw and tile, cells splitting longitudinally, connective Stafleu (1954) are used to indicate the loca­ gland present; style filiform, stigma small; tions of the other specimens: ovary inferior, 2-celled. a with BISH-Bernice P. Bishop Museum, only 1 seed (rarely 2) developing from the Honolulu many ovules, crowned by the persistent calyx GH-Gray Herbarium, Cambridge lobes or by the truncate scars of the calyx K-Royal Botanical Gardens, Kew lobes; umbilicus sometimes present; seeds M - Botanische Staatssammlung, large with a thin membranaceous or a thick Miinchen cartilaginous or fibrous seed coat; cotyledons MICH-University of Michigan, Ann Arbor thick, fleshy, completely free or partly or NY - N ew York Botanical Gardens, entirely fused. New York Type species: L. US-United States National Herbarium, Washington KEY TO THE SPECIES IN HAWAII I have examined the following numbers of A. Flowers in cymes or racemes; calyx tube fruits and seeds of the species included in this extending beyond the ovary. problem. When possible both fresh and dried B. Calyx tube 10-30 mm. long. material was studied. Fresh Dried C. Inflorescence axillary from older Eugenia malaccensis ...... 15 3 leafless nodes or rarely from leafy E. Jambos ...... 5 6 "Calyx tube" is used here, as is customary in E. sandwicensis...... 100 60 treatments of the genus, to refer to the inferior ovary E. Cumini ...... 50 20 and especially to the tissues adhering to i~. There is no E. uniflora ...... 10 5 intention to imply the exact morphologICal n~ture of the inferior ovary of Eugenia by the use of thIS term . E. rariflora ...... 15 The origin of the inferior ovary in this genus has not E. koolauensis...... 20 10 as yet been satisfactorily determined. Eugenia in Hawaii - WILSON 165

nodes; petals not reflexing at an­ verse; leafy branches brown, glabrous, 3-5 thesis; stamens 1-2 cm. long; mm. in diameter, angled or terete; internodes leaves 5-9 cm. wide, elliptical to 1.5-6 cm. long; leaves 14-25 cm. long, 5-9 obovate-oblong ...... cm. wide, elliptical to obovate-oblong, apex ...... 1. E. malaccensis abruptly acute or obtuse, base cuneate to C. Inflorescence terminal; petals re­ abruptly cuneate; margin entire or slightly flexing at anthesis; stamens 2-5 undulate; blade coriaceous, above glabrous, cm. long; leaves 2.5-5 cm. wide, shiny, dark green, sparsely black punctate, linear-Ianceolate .... 2. E. Jambos below pale, glabrous, minutely black punc­ B. Calyx tube 3-8 mm. long. tate; midrib light green, shallowly impressed above, elevated below; primary lateral veins D. Stamens 20-30, included in the alternate or opposite, 8-14 on a side 1-2.5 calyx tube, inflexed, 0.5-1.5 mm. long; fruit pink to deep red ...... cm. apart, irregularly ascending at 140-150°, ...... 3. E. sandwicensis meeting in a conspicuously sinuate, continu­ ous intramarginal vein 0.5-1.5 cm. from leaf D. Stamens more than 50, exserted, margin, smaller, continuous intramarginal spreading, 3-7 mtp. long; fruit vein 1-2 mm. from leaf margin, irregularly dark purple or black. 4. E. Cumini lobed, raised or impressed above, elevated A. Flowers single or in pairs, axillary; calyx below; the veinlets less distinct, raised-reti­ tube not extending beyond the ovary. culate; petioles 1-1.5 cm. long, 3-4 mm. E. Fruits longitudinally 8-ribbed; calyx wide, reddish-green, glabrous; cymes axillary lobes membranaceous .. 5. E. uniflora or on stems below leaves 2-5 cm. long; pe­ E. Fruits not ribbed; calyx lobes fleshy. duncle 5-10 mm. long, angled or terete, glabrous, reddish-green, pedicels when pres­ F .. Leaves flat or slightly concave, ent, 2-8 mm. long, glabrous, reddish-green, glabrous beneath or only sparingly articulate, but some flowers sessile; calyx tube puberulent near midrib ...... green to reddish-green, 1-3 cm. long, 1-1.5 ...... 6. E. rariflora cm. wide, obconic, elevated above ovary, nar­ F. Leaves strongly concave and with rowing into a short pseudostalk 0.5-1.0 cm. the entire lower surface puberulent. long, subtended by 2 deltoid bracts 1-1.5 G. Fruit orange-yellow; flowers mm. long; calyx lobes 4, persistent, broad, with punctiform pistil ...... rounded, 2-3 cm. long, 6-8 cm. wide, green; ...... 7. E. koolauensis petals 4, spreading, obovate-orbicular, pink G. Fruit red; flowers with peltate to red, glabrous, deciduous, glandular-punc­ pistil ...... 8. E. molokaiana tate, 6-9 mm. long, 7-10 mm. wide, apex rounded or acuminate, base truncate; stamens 1. Eugenia malaccensis L., Sp. PI. 470,1753. numerous (about 100), exserted, 1.0-2.0 cm. long; filament slender, red, glabrous, terete Jambosa malaccensis (L.) DC., Pro dr. 3: 286, above, flattened below; anthers white 9-12 1828. mm. long, oblong; style red, subulate, 1.5-2 (L.) Merr. and Perr., Ar­ cm. long, glabrous; ovary 2-celled multi­ nold Arboretum, Jour. 19: 215, 1938. ovulate; fruit obovoid 5-7.5 cm. long, 4-6 8-20 m. tall; branches greyish-brown, cm. in diameter, pinkish to dark red, um­ smooth, glabrous; leaf scars 3-6 mm. wide, bilicate on top, crowned with truncate scars lunate-elliptic, pale; bundle scar large, trans- of the calyx lobes or calyx lobes persisting; 166 PACIFIC SCIENCE, Vol. XI, April, 1957 pericarp crisp, watery, 1.0-2.0 cm. thick; 2. Eugenia Jambos 1., Sp. PI. 470, 1753. seed loosely attached within, subglobose, 1.5-2.0 cm. in diameter; seed coat fibrous, Jambosa Jambos (1.) Millsp., Field Mus. Nat. brown, 1 mm. thick, adhering closely to the Hist., Bot. Ser. 2(1): SO, 1900. rugose surface of the cotyledons; cotyledons (1.) Alston, in lrimen, Fl. 2, white or greenish, equal or unequal, not Ceyl. 6 (Suppl.): 115, 1931. fused. Tree 6-10 m. tall; branches brown to yel­ Common name: "Ohia ai," Mountain lowish-brown, glabrate, longitudinally ridged; . leaf scars rounded shield-shaped, 2.5-4 mm. DISTRIBUTION: Common in the moist wide, pale; stems of leafy branches 3-4 mm. gulches on the larger islands. Native to the in diameter, 4-angled or compressed, becom­ Indo-Malayan region, it may now be found ing terete in age, glabrous; internodes 1.5-3 in cultivation, widely distributed in the tropics cm. long; leaves 10-20 cm. long, 2.5-5 cm. of the world. wide, lanceolate or oblong-lanceolate, taper­ Specimens examined: ing to an acuminate apex, base cuneate; mar­ HAWAIIAN ISLANDS: Hillebrand and Lyd­ gin entire; blade coriaceous, above olive­ gate; Mann and Brigham 119; U. S. Explor. green to reddish-green, glabrous, shiny, Exped. (NY). minutely pustulate, below paler, glabrous, KAUAr: Waioli Valley, along stream, alt. minutely glandular-punctate; midrib shallowly 100 m., Feb. 27, 1927, MacDaniels 909. impressed above, elevated below, light yel­ OAHU: Punaluu, stream bank, elev. SOo ft., lowish-green to reddish-green; primary lateral Sept. 2S, 1930, St. John 10,581 (NY); Punaluu veins alternate or opposite, 10-15 on a side, Valley, in dark forest in wet ground at bot­ 5-15 mm. apart, slightly elevated above, very tom of Pig God trail, Sept. 2, 1932, Degener prominent below, straight or slightly curved 7,349; Waikane-Schofield trail, side of trail, ascending at 140-150°, meeting in an irreg­ 1000 ft. alt., Dec. 2, 1951, Wilson 46; Koolau­ ularly lobed continuous intramarginal vein loa, Kaluanui, Sacred Falls Valley 700 ft. alt., 3-5 mm. from leaf margin; the veinlets ob­ May IS, 1952, Wilson and Doty 137, 138, 139, scure above, distinctly raised-reticulate below; and 141; Kipapa Gulch, frequent at bottom petiole 5-10 mm. long, 2-3 mm. wide, gla­ of gulch, in "koa zone," Egler 37-421; Kalihi brous, dark reddish-green; racemes terminal, Valley, Jan. 1, 1920, Garber 97; Moanalua 6-10 mm. long, rachis 6-15 mm. long, 3-4 Valley, March 7, 1910, Forbes 1465.0; Kaumo­ mm. wide, 4-angled brownish-green to red­ kunui Gulch, rich dark wet gulch at 1500 ft., dish-green, glabrous; pedicels 7-15 mm. April 13, 1936, Degener 11,887 (NY, MICH); long, single on the pedicels, 3-6 cm. Waianae Mts.: Makaleha Valley, Jan. 14, long, 6-s cm. in diameter; calyx tube obconic, 1929, Neal; Puu Kaupakuhale, 2nd gulch of 1-1.5 cm. long, 7-10 mm. wide, elevated N.E. slope of Puu Kaala, in wood, May 14, above ovary, narrowed into a short pseudo­ 1933, St. John 13,173. stalk, glabrous or sparsely puberulous, mi­ MOLOKAr: Mapulehu, April 1910, Rock. nutely glandular-punctate, green or yellow­ MAUl: Kailua, Haleakala, April 1911, Rock. green, subtended by 2 caducous, glabrate, HAWAII: Hilo, May 1909, Faurie. subulate bracts 0.S-1.0 mm. long; calyx lobes Eugenia malaccensis was most likely intro­ 4, persistent, fleshy, unequal, 1 pair 6-S mm. duced into the Hawaiian Islands by the Poly­ long and S-9 mm. wide, the smaller pair 4-6 nesians. It may be found growing in large mm. long, S-9 mm. wide, below minutely groves in moist, shaded valleys where it was glandular-punctate, glabrous or sparsely pu­ probably originally planted and has subse­ berulous; petals 4, white to greenish-white, quently become established. orbicular to ovate-orbicular, concave, spread- Eugenia in Hawaii - WILSON 167 ing, glandular-punctate, 1-1.7 mm. in diam­ initely known from Kauai, Molokai, Oahu, eter, glabrous; stamens numerous (about Maui and Hawaii," no specimens of it were 200), creamy-white, 1-5 cm. long; filament seen from Molokai. slender, terete, creamy-white, glabrous; an­ thers white, oblong, 1-2 mm. long; style 3. Eugenia sandwicensis Gray, U. S. Ex­ terete, subulate, 3-4 cm. long, glabrous, pI or. Exped. Bot. (official ed.) 519, 1854. creamy-white to greenish-white, exserted or (Gray) Ndz., in Engl. included; fruit subglobose, 2-4 cm. tall, 4-6 and Prand pflzfam. 3(7): 85, 1893. cm. wide, yellow or pinkish-yellow, minutely glandular-punctate, umbilicate on top, Eugenia sandwicensis var. parvi/olia Hdb., crowned by persistent calyx lobes, style often FL Hawaii. Is. 129, 1888. persistent; pericarp fleshy, 1-1.5 cm. thick; Syzygium oahuense Deg. and Ludw., Bot. seed loosely attached within, subglobose, Staatsaml. Miinchen, Mitt. 4: 113, 1952. 2-2.5 cm. in diameter; seed coat 1 mm. thick, Tree or , 3-25 m. tall; branches grey­ brown, coriaceous, closely adhering to the ish-brown to reddish-brown, glabrous; leaf surface of the cotyledons; cotyledons white scars 1-4 mm. wide, rounded, shield-shaped, or greenish-white, equal or unequal, not reddish-brown to yellowish-brown; young fused. leafy branches green to reddish-green, gla­ Common name: "Ohia loke," Rose Apple. brous, 1-4 mm. in diameter, distinctly 4- DISTRIBUTION: Sparingly naturalized on angled, angles winged; wings 0.2-2.0 mm. probably all of the larger islands of Hawaii. wide, branchlets becoming terete with age, Widely distributed in the tropics of the world. dark red to reddish-brown bark scaling off in Specimens examined: longitudinal strips exposing yellow-grey to KAUAI: Kokee Camp, becoming natural­ reddish-yellow bark beneath; internodes 1-5 ized,July 5,1926, Degener 7,341 (NY); Waioli cm. long; leaves 2-14 cm. long, 1.5-5.0 cm. Valley, along stream, alt. 50 m., Feb. 27, 1927, wide, obovate, ovate, elliptic or ovate-Ianceo­ MacDaniels 908. late, apex acute, obtuse, retuse, or apiculate, base truncate to cuneate, blade coriaceous, OAHU: Waikane-Schofield trail, Waikane, flattened or concave, margin entire, slightly 750 ft. alt., side of road, Dec. 2, 1951, Wilson revolute (rarely strongly so), above dark green 45; Waiahole, Jan. 23, 1909, Rock 1,285 and or yellowish-green, shiny, glabrous, minutely 1,287; Manoa, near Woodlawn, spreading glandular-punctate, below paler, dull, gla­ locally, Apr. 1937, Egler 37-423. brous, minutely glandular-punctate, midrib MAUl: lao Valley, Wailuku, roadside, elev. pink to dark red, shallowly impressed above, 800 ft., Feb. 9, 1930, St. John 10,277 (BISH, elevated below; primary lateral veins alternate NY). or opposite, 15-30 on a side, 2-8 mm. apart, HAWAII: Naturalized, Degener 7,344; South irregularly ascending at 100-115°, meeting in Kona, Honomalina, Ranch House, near Kona an irregularly lobed intramarginal vein 0.5- highway, 1800 ft. alt., Sept. 7, 1952, Chock 1.5 mm. from leaf margin, raised on both 768; near Glenwood, naturalized in pasture, surfaces but more distinct below; the veinlets June 23, 1929, Degener 7,343 (NY). raised-reticulate; petioles 2-10 mm. long, 1-2 Degener (1932-34) records that Eugenia mm. wide, reddish-brown, glabrous, cymes Jambos was probably first introduced into simple or compound, in axils of upper leaves, Hilo from Rio de Janeiro by Mr. Bridge in 5-8 cm. long; peduncle 2.5-3.5 cm. long, 1853. It may now be found growing along 1.5-3 mm. wide, 4-angled, winged, yellow­ road and trail sides and in other moist areas. green to reddish-green, pedicels 2-4 mm. Although Degener also records it as "def- long, articulate; calyx tube turbinate, 3-4 168 PACIFIC SCIENCE, Vol. XI, April, 1957 mm. long, 3.5-5 mm. wide, glabrous, reddish­ Lydgate; Wahiawa Mts., August 1909, Forbes green to yellow-green, minutely glandular­ 181.K; Wahiawa, Kahili Swamp, 2100 ft. alt., punctate, subtended by two deciduous, Lihue-Koloa Forest Reserve, Dec. 29, 1930, glabrous, deltoid bracts 1-2 mm. long; the St. John et al. 10,850; Koloa, Laaukahi ridge, 4 calyx lobes 0.5 mm. long, imbricate, ob­ % mile north ofN.W. facing slope, 850 ft. alt., tuse, reddish-green to dark red, early decid­ moist wooded gulch, Dec. 24, 1947, St. John, uous; petals 4, white to greenish-white, Webster and Wilbur 23,007; Laaukahi, Haiku, spreading, soon deciduous, usually discrete 1300 ft. alt., dense woods on precipitous but sometimes united and falling off as a slope, Dec. 22, 1933, St. John and Fosberg calyptra, ovate or obovate, often emarginate, 13,486; Ka Loko Reservoir (Kilauea), Oct. 8, glabrous, glandular-punctate, apex subacute 1916, Forbes 544.K; E. fork of Kilauea River, or obtuse, base truncate, 2-3 mm. long, 2-3 rain forest, alt. 400 m., Feb. 11, 1927, Mac­ mm. wide; stamens (about 30) inserted on the Daniels 654; Wainiha, Wainiha Valley, moist margin of the calyx tube, 0.5-1.5 mm. long, lower forest, 800 ft. alt.,Jan. 1, 1934, St. John introrse, included; filaments white to pinkish, and Fosberg 13,929; Wainiha, Wainiha Valley, subulate, glabrous; anthers white 0.4-0.8 mm. 1000 ft. alt., on bank, side of road, in native long, orbicular-ovate; pistil white to reddish, forest, Dec. 31, 1952, Wilson 235; Hanakapiai, glabrous, slender, 0.8-1.5 mm. long, in­ Napali Coast, forest on cliff, Jan. 2, 1931, St. cluded; ovules 4-10 in a cell; fruit light pink John et al. 10,992; Hanakoa, Waiahuakua to dark red, 4-10 mm. high, 5-10 mm. wide, Stream, 350 ft. alt., Dec. 31, 1952, Wilson and glabrous, sqiny, minutely glandular-punctate, St. John 238; Hii Mts., Oct. 22, 1916, Forbes globose or elliptic, flattened on top, crowned 652.K. by truncate scars of the calyx lobes; pericarp OAHU: No locality: Mann and Brigham 204; fleshy, 1-1.5 mm. thick; seed globose, elliptic Hillebrand (received July 1865) 311 (GH) or oblong-elliptic; seed coat reddish-brown, (locality illegible, probably Oahu); U. S. 0.5 mm. thick, loosely adhering to the peri­ Explor. Exped. (US). Koolau Mts.: Kahuku carp, closely adhering to the smooth surface Army trail, July 1930, Russ; Hauula, Kaipapau of the cotyledons; cotyledons equal, greenish Forest Reserve, Maakua-Papali ridge, wooded or white, conspicuously glandular-punctate, ridge, 1200 ft. alt., St. John 13,372; Kahana not fused. Valley, head of, Hauula Forest Reserve, 1000 Type: U. S. Explor. Exped. "Oahu, Sand­ ft. alt., lower woods, Dec. 10, 1933, St. John wich Islands; on the mountains behind Ho­ 13,410; Kahana Valley (south side of), ridge nolulu" (US). mauka of church, dense forest at 1500 ft., Common name: "Ohia ha," known on Nov. 5, 1950, Degener and Silva 21,069 (US); Maui as "Paihi." Kahana Valley, head of, 1500 ft. alt., Aug. DISTRIBUTION: Endemic to the Hawaiian 31, 1924, Harris; Kaipapau on S. slope of Islands. Found in the moist forests on Kauai, ridge, 2500 ft. alt., Oct. 15, 1933, Suehiro; Oahu, Molokai, Lanai, and Maui; not known Punaluu, Dec. 3-4, 1908, Rock 632 and 687 from the island of Hawaii. (GH), and 526; Punaluu, Dec. 24-29, 1908, Specimens examined: Rock 377 (GH); Punaluu, Dec. 3, 1908, Rock KAUAr: Waimea, Alakai Swamp trail, 3800 149 (GH); Punaluu, wet mountain side, Nov. ft. alt., rain forest, Dec. 25, 1952, Wilson 206; 30, 1929, Tanaka; Punaluu to Kaipapau, May on Kaholuamanu above Waimea, Sept. 2-9, 8-13, 1909, Forbes; Punaluu to Kaipapau, 1895, Heller 2,241; Kaholuamanu to Waimea, May 3-8, 1909, Forbes and Cooke, and Forbes Oct. 27, 1916, Hitchcock 15,558 (US); near and Thompson; Punaluu to Kaipapau, Nov. Kaholuamanu, Kauluwehi Swamp, Oct. 25, 14-21, 1908, Forbes; Kahana, Kaluanui, open 1916, Hitchcock 15,520 (US); Wahiawa Mts., woods, 2000 ft. alt., Nov. 30, 1929, St. John Eugenia in Hawaii - WILSON 169

10,099 (US); Waikane-Schofield trail, Wai­ 1, 1926, MacDaniels 98; Pauoa, Konahuanui kane, 750-1250 ft. alt., Oct. 16, 1932, Krauss; trail, Feb. 15, 1921, Garber 229; Pauoa flats, Waikane-Schofield trail, near summit, 2000- 1926, Skottsberg 1,783; Manoa, slopes back 3000 ft. alt., Sept. 16, 1932, Yuncker 3,186 of Woodlawn, on crest of ridge, alt. 1000 ftc, (US); Waikane-Schofield trail, 2200 ft. alt., Dec. 31, 1942, Kuykendall 49; Olympus trail, Dec. 2, 1951, Wilson 44; north ridge of Kaa­ alt. 700 m., Dec. 21, 1926, MacDaniels 125; awa Valley, April 12, 1931, St. John 11,085; Palolo-Manoa ridge, alt. 350 m., Dec. 21, Heeia, Haiku Valley, Waiahole Forest Re­ 1926, MacDaniels 122; Palolo Valley, near serve, 500 ft. alt., in Dicranopteris thicket on first falls of Palolo, Nov. 16, 1919, Garber 70; ridge, Dec. 11, 1932, St. John 12,260; Kona­ Palolo Valley, Oct. 22, 1914, Forbes 1929.0; huanui, Jan. 16, 1909, Forbes 1,040; Kona­ ridge between Palolo and Waialae-iki, Jan. huanui-Forbes 1,003, and 1,309, Bryan 208, 30, 1917, Forbes 2411.0; Palolo-Waialae ridge, MacDaniels 124, and Heller 2,241 (US, MICH); Jan. 27, 1927, MacDaniels 484 and 487; Wili­ Pupukea Military trail, Jan. 29, 1927, Mac­ wilinui ridge, 1600 ft. alt., March 16, 1952, Daniels 549; Waimea-Malaekahana, 1900 ft. Wilson 106, 108, 111, 113, 115, 116, 117, 118, alt., March 22, 1953,Ozaki 391,392,390, and 119, and 120; Niu Valley, summit ridge, Aug. 389; Paalaa, South Opaeula Gulch, Nov. 9, 22, 1909, Rock 4,840. Waianae Mts.: Dupont 1930, St. John 10,630; Waipio, Kipapa Gulch, trail, northern slope of Kaala, Feb. 29, 1949, wet high mountain ridge, Nov. 10, 1929, Degener et at. 19,428; Dupont trail, north Tanaka; Waipio, Kipapa Gulch, E. of Puu slope of Mt. Kaala, 2800 ft. alt., rain forest, Kamana, wooded ridge, 1700 ft. alt., May 15, Sept. 10, 1950, Hatheway et al. 344; eastern 1932, St. John 11,683; Waipio, Kipapa Gulch, part of Kaala summit, Sept. 25, 1938, Degener south ridge, wooded slope, 1300 ft. alt., Nov. et al. 12,247 (NY); Kaala, Oct. 13, 1929, 10, 1929, St. John 10,037; Waipio, Kipapa Yoshinaga; Mokuleia, 1200 ft. alt., July 18, Gulch, south ridge, 1400 ft. alt., woods, 1924, Wilder 1,287; Makaha Valley, Feb. 12- Oct. 29, 1929, St. John 9,965; Kawailoa, Ka­ 19, 1909, Forbes; Honouliuli, ridge above Ku­ waiiki Ditch trail, alt. 1340-1000 ft., Nov. 2, pehau, dry brushy slope, alt. 650 m., June 1947, Wilbur 257 (US); Kawailoa, Kawaiiki 30, 1935, Fosberg 10,986; Palehua, Aug. 23, Ditch trail, moist forest, 1050 ft. 1922, Skottsberg 207; slope south of Palawai alt., Jan. 17, 1953, St. John 24,973; Kawailoa, Gulch, ca. 2700 ft., March 27, 1948, Wilbur Kawaiiki Ditch trail, Aug. 15, 1922, Skottsberg 602. 230; Kawailoa trail, rain forest, Oct. 31, 1937, MOLOKAI: Olokui, 3000 ft. alt., ridge be­ Hartt; Kalauao ridge, Ewa Forest Reserve, tween upper forks of Wailele stream, rain moist woods, March 29, 1933, St. John forest, Feb. 6, 1948, St. John and Wilbur 13,028; Halawa ridge trail, in moist forest, 23,343; Halawa, ridge south of valley, Aug. April 25, 1948, Cowan 973; mauka of Red 1912, Forbes 477.Mo. (US); Kaluaaha, April Hill, in forest, Oct. 9, 1932, Degener 7,328 1910, Rock 7,060 (GH); Kaluaaha, rain forest, (US, MICH); Kalihi-Nuuanu ridge, alt. 550 June 28, 1928, Degener 7,323 (GH); Pukoo, m., Jan. 23, 1927, MacDaniels 156; Kalihi­ June 1910, Faurie 434. Nuuanu, Puu Lanihuli, main ridge running LANAI: Aug. 1913, Munro 15; Sept. 1917, S.W. from Kalihi-Nuuanu, wooded ridge, Forbes 361.L (US); Waiakeahua Gulch, in de­ 1600 ft. alt., Nov. 29, 1931, St. John 11,176; cadent, deer devastated forest at 2500 ft., Kalihi-Nuuanu, Lanihuli trail, Sept. 17, 1908, Aug. 4, 1949, Degener and Murashige 20,322 Forbes; Lanihuli trail, Nov. 14, 1908, Forbes; (US); Waiakiola Valley, July 28, 1910, Rock Lanihuli trail, Dec. 10, 1908, Forbes; Nuuanu­ 8,056 (GH): Kaiholena, Munro 6; Lanaihale, Kalihi ridge, Aug. 13, 1922, Skottsberg 156; July 28, 1940, Degener 12,976 (US); Puu Aalii, slope mauka from Tantalus, alt. 600 m., Nov. Kealia Aupu-Kaunolu divide, lower forest, 170 PACIFIC SCIENCE, Vol. XI, April, 1957

April 14, 1938, St. John 18,851; mountains Tree 6 to 20 m. tall; branches pale yellow­ near Koele, June 1913, Forbes 78.L; moun­ ish-grey, glabrous, in age the bark greyish­ tains near E. end, June 1913, Forbes 275.L. white; leaf scars 2-4 mm. wide, rounded MAUl: Makawao, Oct. 1910, Rock 8,617 shield-shaped, yellowish-grey; leafy branch­ (US); April 1911, Curran 62 (US); upper lets 1-3 mm. in diameter, terete or slightly ditch trail Haleakala, Oct. 13, 1922, Skottsberg angled, glabrous, pale brown to greyish­ 808; Kailua, north slope of Haleakala, June white; internodes 1.5-4.5 cm. long; leaves 13, 1920, Forbes 2499. M (US); Kailua, Dec. opposite, 7-18 cm. long, 3-8 cm. broad, 25, 1908, Rock; Kipahulu, 2500 ft. alt., Kau­ oblong-ovate to elliptic-oblong, apex shortly kaua Gulch, west ridge, summit of, Acacia or abruptly acuminate, rarely obtuse, base Koa woods, Dec. 28, 1936, St. John and Catto broadly cuneate narrowing toward petiole, 17,806; Nahiku, July 1910, Forbes; along margin entire or slightly undulate, blade cori­ ditch near Oopuola stream, lower rain forest, aceous, above olive-green, minutely glandu­ July 7,1927, Degener 7,324 (US); Puu Kukui, lar-punctate, glabrous and shiny, below dark upper forest, 3-5000 ft., Sept. 25, 1916, yellowish-green, dull, minutely pitted or Hitchcock 14,804 (US); near where trail leaves pustulate, glabrous; midrib shallowly im­ tunneled stream for Mt. Eke climb, rain pressed above, elevated below; primary lateral forest, Aug. 27, 1927, Degener and Wiebke veins numerous, alternate or opposite, 25-40 2,307 (US, MICH). on a side, 2-5 mm. apart, irregularly ascending This species exhibits extreme variation, at 140-150°, meeting in an irregularly lobed especially in the leaves. The leaves are of intramarginal vein 1-3 mm. from leaf margin, various shapes, with a great number of inter­ raised on both surfaces; the veinlets raised­ mediate forms, and they range in size from reticulate; petioles 1-2.5 mm. long, 1 mm. 14 X 5 cm. to 2 X 1.5 cm. The obovate leaf wide, glabrous; inflorescence cymose, on pre­ with an obtuse apex is most frequently en­ vious years' branches or occasionally in axils countered, but even on a single branch with of the leaves, rarely terminal, 4-12 cm. long; leaves predominantly of this shape, others peduncle 1-3 cm. long, terete or slightly an­ with ovate leaves and acute apices may be gled, glabrous, subtended by two deciduous, found. Of all the characters the flowers are subulate bracts 0.5-1.5 mm. long, glabrous; the most constant. pedicels decussate, articulate, slender terete No combination of characters was found or slightly angled, subtended by one decidu­ on which a separation of the various forms ous subulate bract; flower sessile, calyx tube could be based. Any attempt at reclassifying campanulate, 3-5 mm. long, 2-3 mm. across, the group always disclosed intergrading forms brownish-pink, glabrous, finely glandular­ which rendered the system impractical. punctate, narrowing into a stout psuedostipe, The absence of this widely distributed spe­ subtended by 2 deciduous bracts; bracts cies on the island of Hawaii is peculiar. Rock reddish-brown, deltoid, 1-1.5 mm. long; (1913a) lists this species as occurring "on all calyx lobes 4, quickly deciduous; petals 4, islands of the group," but no single collection white, minutely glandular-punctate, orbicular, of it was found from Hawaii. concave,2-3 mm. in diameter, falling off as a calyptra; stamens numerous (about 100), ex­ 4. Eugenia Cumini (L.) Druce, Bot. Exch. serted and spreading, inserted on the margin Club Brit. Is., Rpt. 3: 418, 1914. of the calyx tube; filaments pinkish, 2-6 mm. Cumini L., Sp. PI. 471, 1753. long, slender, subulate, glabrous, finely Eugenia Jambolana Lam., Encycl. 3: 198, 1789. glandular-punctate, anthers white, 3-7 mm. (L.) Skeels, U. S. Dept. Agr., long, orbicular-ovate; style terete, subulate, Bul. 248: 25, 1912. included, 5-6.5 mm. long, glabrous, finely Eugenia in Hawaii - WILSON 171

glandular-punctate; berry oblong to oblong­ along bank of stream, Dec. 29, 1951, Wilson elliptic, asymmetric 1.5-2.0 cm.long, 1.0-1.5 104. cm. wide, glabrous, dark purple or black, HAWAII: Puna, Pahau Nui, alt. about 1700 shiny, minutely glandular-punctate, umbili­ ft. associated with guava, Aug. 2, 1945, cate, crowned by the truncate scars of the Fagerlund and Mitchell 1,104; above Hilo, calyx lobes, umbilicus 1-2 mm. tall, 1.5-3 along Malili stream, homestead, alt. 650 m., mm. in diameter; pericarp pulpy, 1.5-3 mm. MacDaniels 249. thick, seed ellipsoid or oblong-ellipsoid, 1.0- Eugenia Cumini is of recent introduction 1.5 cm. long, 0.5-1.0 cm. wide, seed coat, into the Hawaiian Islands. It has rapidly be­ crustaceous, 0.5-1.0 mm. thick, closely ad­ come established and widely distributed, most hering to the subrugose surface of the coty­ likely by birds which relish the fruit. It may ledons; cotyledons unequal, not fused, con­ now be found in abundant stands in the dryer spicuously glandular-punctate. regions of the islands, forming the dominant Common name: Java Plum. vegetation of valleys with periodically dry DISTRIBUTION: Widely distributed on all streams. the Hawaiian islands, found in large stands in No record has been found that indicates dry and moist valleys; cultivated, spreading when this species was introduced into the from cultivation, and established. Widely dis­ islands. The earliest collection of it .in the tributed in the Indo-Malayan region and in Hawaiian Islands was by Faurie on Maui in the tropics of the world. August 1909. He records it as "Wailuku Specimens examined: Culta." KAUAr: Hanapepe, Koula Valley, 750 ft. alt., on side of road, Dec. 27, 1952, Wilson 5. Eugenia uniflora 1., Sp. PI. 470, 1753. 210. Shrub 2-3 m. tall, branches greyish-brown, OAHU: Koolau Mts.: Honolulu, Nuuanu smooth, glabrous; leaf scars 1-1.5 mm. wide, Valley, Dowsett Highlands, by road, Sept. 19, rounded shield-shaped, yellow-brown to red­ 1943, Neal. Waianae Range: Kamananui, Du­ dish-brown; leafy branches green to greyish­ pont trail on ridge south of Pamoa Gulch, by brown, glabrous, 1-4 mm. in diameter, angled forest reserve fence, 1500 ft. alt., Sept. 14, when young, becoming terete in age; inter­ 1952, Wilson 145; N. of Kaala, naturalized in nodes,2-4 cm. long; leaves 4-7 cm.long, 2-4 pasture, April 26, 1937, Degener 11,902 and cm. wide, ovate to ovate-Ianceolate, tapering 11,903 (NY); Mokuleia, Makaleha Valley, to acuminate apex, base rounded to sub­ 900 ft. alt., in lowland scrub, Sept. 15, 1950, cuneate; margin entire, slightly revolute; Hatheway et al. 357; Honouliuli, Puu Mana­ blade thin coriaceous, above glabrous, shiny wahua, lower woods, 1800 ft. alt., Sept. 29, yellow-green to dark olive-green, minutely 1929, St. John 9,893. glandular-punctate, below pale yellow-green, MOLoKAr: Halawa Valley, 150 ft. alt., glabrous or very sparsely puberulous, mi­ spreading along trail, by abandoned, over­ nutely glandular-punctate; midrib reddish­ grown taro patches, Dec. 27, 1932, St. John green, shallowly impressed above, elevated et al. 12,676; eastern side of Wailau Valley below; primary lateral veins alternate or op­ near ocean, naturalized, Aug. 14, 1928, Deg­ posite, elevated on both surfaces but more ener 9,664 (NY); near Kanalo, naturalized, conspicuous below, 9-12 on a side, 4-7 mm. Aug. 8, 1928, Degener 7,345 (NY). apart, irregularly ascending at 140-150°, meet­ MAUl: Wailuku, culta, Aug. 1909, Faurie ing in an irregularly lobed continuous mar­ 55; Muolea, E. Maui, 3 miles from Hana, ginal vein 2-5 mm. from leaf margin; less along roadside, Dec. 27, 1951, Wilson 101; conspicuous continuous or sometimes inter­ Ukumehame, Ukumehame Gulch, 800 ft. alt., rupted marginal vein 0.5-1 mm. from leaf 172 PACIFIC SCIENCE, Vol. XI, April, 1957

margin; the veinlets less distinct, raised­ 6. Eugenia rariflora Benth., in Hooker's reticulate; petioles 2-5 mm. long, 0.5-1 mm. London Jour. Bot. 2: 221, 1843. wide, dark reddish-green, glabrous; flowers Eugenia waianensis Deg., Fl. Hawaii., Fam. single or in pairs, axillary; peduncles 2-4 cm. 273, 7/15/32. long, 4-angled or terete, glabrous; calyx tube 2-3 mm. long, 3-4 mm. wide, green, glabrous Deg. var. glabra Deg., Fl. or sparsely puberulent; calyx lobes 4, per­ Hawaii., Fam. 273, 8/10/32. sistent, 3-4 mm. long, 3-4 mm. wide, mem­ Eugenia rariflora Benth. var. parvifolia Hdb., branaceous, green, ovate with acute apex, Fl. Hawaii. Is. 129, 1888. ciliate, minutely glandular-punctate, attached Tree or shrub 3-7 m. tall; branches grey to to annular disk within the tube; petals 4, greyish-white, glabrous, in age the bark grey­ white, spreading, persistent, obovate with an ish-white to greyish-brown, longitudinally obtuse apex, ciliate, 7-8 mm. long, 5-6 mm. and transversely irregularly furrowed; leafy wide, sparsely glandular-punctate; annular branchlets 1.5-2 mm. in diameter, quadran­ disk raised 0.5 mm. above point of petal in­ gular to terete, brown, brown puberulent sertion, strigose; stamens numerous (about when young; leaf scars rounded shield-shaped, 50), spreading, inserted on annular disk; fila­ 1-1.5 mm. wide; bundle scar 1; internodes ments 3-7 mm. long, white glabrous, sub­ 0.5-3 cm. long; leaves opposite, 3-7 cm.long, ulate; anthers white, 0.5-1 mm. long, orbic­ 1.5-5 cm. wide, blade elliptical, ovate or ular-ovate; style, terete, subulate, slightly obovate, apex obtuse, acute, retuse or apic­ exserted or included, 4-6 mm.long, glabrous; ulate, base cuneate or rounded, coriaceous, fruit subspherical, red, 1-2 cm. high, 1-2 cm. above olive-green, shiny, minutely glandular­ wide, glabrous, minutely glandular-punctate, punctate, glabrous or sparsely puberulent crowned by the persistent calyx lobes, con­ near midrib, below dull, yellowish-green, spicuously longitudinally 8-ribbed; pericarp minutely pustulate, glabrous or somewhat fleshy, 2-3 mm. thick; seed subglobose, 8-15 puberulent at base near midrib; midrib shal­ mm. in diameter, seed coat thin membranous, lowly impressed above, elevated below; pri­ looseiy adhering to the pericarp or to the mary lateral veins alternate or opposite, 5-3 surface of the cotyledons; cotyledons fused, on a side, 4-8 mm. apart, irregularly or finely glandular-punctate. straight ascending at 140-150°, meeting in an Common name: Surinam Cherry, Pitanga. irregularly lobed intramarginal vein, 1-2 mm. DISTRIBUTION: Very sparingly naturalized from leaf margin, raised on both surfaces, but on the larger islands. Native to . more distinct below; the veinlets raised-retic­ Specimens examined: ulate; petioles 2-4 mm. long, 1 mm. wide, KAUAI: Lihue, planted, June 10, 1926, brown, glabrous or slightly puberulent; Degener 2,080. flowers single or in pairs, axillary; peduncles OAHU: U. S. Expt. Station, Oct. 25, 1926, 5-1'5 mm. long, glabrous or puberulent; calyx MacDaniels 349. tube 2-3 mm. long, 3-4 mm. across, obconic, MOLOKAI: Halawa Valley, 50 ft. alt., shrubs brown, puberulent, minutely glandular-punc­ by roadside, Dec. 27, 1932, St. John et al. tate, subtended by two persistent, subulate 12,665. sparsely puberulent bracts 1-1.5 mm. long; HAWAII: Planted and persisting, April 10, calyx lobes 4, unequal in length, 1 opposite 1930, Degener 7,331 (NY). pair 3-4 mm. long, 3-4 mm. wide, the other The number of collections of this species is pair shorter, 2-3 mm. long, 3-4 mm. wide, very meager, and future collections should be ovate, green, fleshy, persistent, below mi­ made to establish the occurrence of it in the nutely glandular-punctate, glabrous, above Hawaiian Islands. glabrous, within the tube attached to annular Eugenia in Hawaii - WILSON 173 disk; petals 4, white, spreading, persistent, ft. alt., steep talus slope, in dryland forest, 6-s mm. long, 4-5 mm. wide, inserted on the. Aug. 31, 1950, Hatheway 376; Mokuleia, east margin of the disk, ovate, obovate or elliptic, side of east branch of Makaleha Valley, minutely glandular-punctate, membranace­ densely forested slope at 1600 ft.,July 2, 1950, ous, ciliate and sparsely puberulous, apex ob­ Degener, Hatheway and Greenwell 20,824 (US); tuse or acuminate, base truncate; annular disk Mokuleia, west branch of E. Makaleha Val­ raised 0.5 mm. above point of petal insertion, ley, lS00 ft. alt., dry forest on steep valley glabrous; stamens numerous (about 150), in­ side, June 11, 1952, St. John 24,827; Moku­ serted on disk; filaments white, 1.5-5 mm. leia, west branch, E. Makaleha Valley, in small long, slender, subulate, glabrous; anthers side gulch, 1750 ft. alt., Sept. 30, 1950, white, 0.5-0.S mm. long, orbicular-ovate; Hatheway 384; Makaleha Valley, May 2, 1915, style terete, subulate, slightly exserted or in­ Rock 17,005 (GH); Makaleha Valley, Oct. 23, cluded, 2-4 mm. long, glabrous; fruit ovoid 1936, Meebold (Degener 21,980) (M); Moku­ to subspherical, crowned by persistent calyx leia, gulch southwest of Dillingham Ranch, lobes, 1-2 cm. high, 1-1.6 cm. wide, glabrous in shade at 1700 ft., April 23, 1950, Degener, or somewhatappressed-pilosulous, red, orange Hatheway and Carrol 20,615; Mokuleia, 4th or yellow, minutely glandular-punctate; peri­ gulch east of Puu Kaupakuhale, Kamananui, carp fleshy, 1-2 mm. thick, seed globose, Puu Kaala, Oct. 23, 1932, Yuncker and Hosaka S-lO mm. in diameter; seed coat thin mem­ 3,216 (US); half mile southwest of Pohakea branous, closely adhering to the pericarp; Pass, single, dying tree on dry grass and cotyledons 2, white or yellowish, minutely lantana covered slope, July 30, 1932, Degener glandular-punctate, usually fused, but some­ and Bush 4,194 (GH); Honouliuli, ridge above times only partly fused or entirely free. Kupehau, dry lantana covered slope, June 30, Lectotype: "Feejee Islands" (Fiji) Hinds 1935, Fosberg 10,995; Honouliuli, Kanehoa 1841 (K). Gulch, dry slope, elev. 700 m., Oct. 12, 1927, Common name: "Nioi" (Rock, 1913b). Judd 65; Honouliuli, between Palehua and DISTRIBUTION: High islands of Polynesia. Palikea, near summit ridge, Dec. 16, 1935, In the Hawaiian Islands it is found on all the Degener et al. 11 ,300; third small valley north­ large islands except the island of Hawaii. east of Palikea (contains pipe line arising Specimens examined: from tunnel), dry woods, Sept. 19, 1932, KAUAI: Hanakoa, 750 ft., Aug. 2S, 1926, Degener, Park and Bush 7,303 (GH, MICH, Judd 44; Haupu, Kipu, soo ft. alt., wooded NY); north of Puu Pane, sunny gulch at 200 slope, Dec. 25, 1933, St. John and Fosberg ft. elev., south of ruin of cathedral, March 26, 13,616; northeast of Kipu, June 17, 1926, 1950, Degener and Carroll 20,581 (US); on Degener 7,304 (NY). firebreak trail 1 m. north of Puu Kaua, 1700 OAHU: Koolau Mts.: Waimano, 6-1919, ft. alt., on dry slope, Sept. 5, 1952, Wilson, Russ; Waimano, E. branch, E. slope above St. John and St. John 143; on firebreak trail, pass, May 29, 1933, Russ; Waimano, Oct. 1 m. N. of Puu Kaua, alt. 1700 ft. on dry 1935, Meebold (Degener 20,768) (M); ridge slope, Feb. 29, 1945, Cowan 843; Kahanahaiki, between Niu and Wailupe, April 11, 1917, 234 m. alt., dry river bottom, Oct. 16, 1925, Forbes 2458.0. Waianae Mts.: Near Kawaiha­ Judd 23; southern slope of Kahanahaiki Val­ pai, shaded dry slope, Jan. 27, 1929, Degener ley, dry forest, Nov. 1, 1931, Degener et al. and Bush 7,305 (GH, NY); Mokuleia, Kaa­ 7,296 (GH, MICH, NY); Makua Valley, awa (= Kaawa?) Gulch, north of Kaala, May 1930, Russ; small gulch on south side of dryish forest slope, Aug. 2, 1935, Degener and upper Makua Valley, dry forest, May 10, Odonez 12,192 (US, NY); Mokuleia, west side 1931, Degener, Park and Bush 7,295 (GH, of east branch of E. Makaleha stream, 1300 NY); Waianae, Feb. 1930, Meebold 8,62~ 174 PACIFIC SCIENCE, Vol. XI, April, 1957

(M); U. S. Explor., Exped., highland near 7. Eugenia koolauensis Deg., Fl. Hawaii. Waianae; large branch of Lualualei Valley, Fam. 273, 8/10/32. southwest of Pohakea Pass, Aug. 4, 1932, Tree 4-7 m. tall; branches grey, glabrate, Degener and Bush 7,299 (NY). in age the bark reddish-brown, black spotted; No locality: Collect. Dr. Hillebrand, Oahu, leaf scars 1-2 mm. wide, rounded shield­ Maui (GH). shaped, reddish-brown; leafy branchlets 1-3 MOLOKAI: Central Molokai, wet forest, mm. in diameter, 4-angled to terete, densely Oct. 13, 1916, Hitchcock 15,193 (US). brown subappressed-pilosulous; internodes MAUl: Olowalu Valley, May 19, 1920, 9-19 mm. long; leaves 2.5-5 cm. long, 1-3.3 Forbes 2417.M (US); Wailuku, W. Maui, Hil­ cm. wide, obovate to elliptic, apex obtuse or lebrand and Lydgate. apiculate, base subcuneate, blade coriaceous, Eugenia rariflora was first recorded for the concave, margin entire, more or less strongly Hawaiian Islands by Hillebrand in 1888. It revolute, above olive-green, minutely glandu­ occurs occasionally in the drier regions of the lar-punctate, glabrous and shiny or subap­ larger islands except Hawaii. This species pressed-pilosulous near veins with inter­ shows considerable variation in its leaves al­ vals glabrous, below pale yellowish-green, though not to as great a degree as E. sand­ minutely pitted or pustulate, subappressed­ WlcenS1S. puberulent; midrib shallowly impressed above, Both E. waianensis and E. koolauensis var. elevated below; primary lateral veins, alter­ glabra are here reduced to Eugenia rariflora nate or opposite, 5-7 on a side, 5-10 mm. because of intermediate forms which make it apart, irregularly ascending at 140-150°, impossible to set definite limits upon the two. meeting in an irregularly lobed intramarginal The concave leaves would be the only char­ vein 1-2 mm. from leaf margin, raised on acter which would separate E. koolauensis var. both surfaces but more distinct below, rarely glabra from E. rariflora. But a collection from slightly shallowly impressed above; the vein­ Palikea, Oahu (Degener 7,303) has leaves lets raised-reticulate; petioles 2-4 mm. long, which are only slightly concave; another spec­ 1 mm. wide, brown subappressed-pilosulous; imen (Degener 20,581) shows leaves that are flowers single or in pairs, axillary; peduncles more strongly concave. This character is cer­ 1-8 mm.long, subappressed-pilosulous, brac­ tainly not a reliable one, and is of no taxo­ teate; bract subulate, yellow-brown subap­ nomic value in this case. This conclusion is pressed-pilosulous, 1 mm. long, 1 mm. wide; supported by the fact that the characters of calyx tube obconic, 2-3 mm. long, 3-4 mm. E. koolauensis var. glabra are identical in all across, subappressed-pilosulous, sub tended other respects with those of E. rariflora. by two persistent subulate brown subap­ The size of the leaves of E. rariflora might pressed-pilosul04S bracts 1-1.5 mm. long; seem to offer a remarkably valuable taxonomic calyx lobes 4, of unequal length, 1 opposite character, particularly when the extremes in pair 3-4 mm. long, 3-4 mm. wide, the other form are considered. E. waianensis is described pair 2-3 mm. long, 3-4 mm. wide, ovate, as differing from E. rariflora in having smaller green, fleshy, persistent, below finely yellow leaves. This character seems to be of value glandular-punctate, appressed-puberulous, when specimens with extremely small leaves above glabrous, within the tube attached to are studied (Degener 7,296). However, some annular disk; petals 4, white, spreading, per­ specimens, such as those collected by Forbes sistent, inserted on margin of disk, concave, (2417.M) and Rock (17,005), have leaves ovate or obovate or elliptic, apex subacute or which show a series of intermediate sizes be­ obtuse, base truncate, 6-8 mm. long, 4-5 tween E. waianensis and E. rariflora. In such mm. broad, membranaceous, minutely glan­ cases this character proves to be of little value. dular-punctate, ciliate and sparsely puberul- Eugenia in Hawaii - WILSON 175

ous; annular disk raised 0.5 mm. above point distinctly concave leaves which have sub­ of petal insertion, puberulent; stamens nu­ appressed puberulence on the lower laminar merous (about 150), inserted on disk occupy­ surface. It is a rare species and occurs in the ing 0.5 mm. of the marginal region; filaments rain forest of the Koolau Range on Oahu. white, 1.5-5 mm. long, slender, subulate; anthers white, 0.5-1.0 mm. long, orbicular­ 8. Eugenia molokaiana K. Wilson and J. ovate; style terete, subulate, slightly exserted F. Rock, sp. nov. or included, 2-4 mm. long, sparsely pilose; ovules 8 in a cell, berry ovoid and asymme­ Figs. 1 and 2 tric, 1.2-2.0 cm. high, 1.2-1.6 cm. wide, Arbor 2.5-3 m. alta, ramulis folios is 0.5-1.5 sparingly appressed-pilosulous, light orange­ mm. diametro manifeste tetragonis usque yellow, shiny, minutely yellow glandular­ teretibus, crebrissime brunneis subadpressi­ punctate, crowned by the persistent calyx pilosulis, internodis 7-24 mm. longis, foliis lobes; pericatp fleshy, 1-2 mm. thick; seed oppositis 2.0-3.0 cm. longis, 1.4-2.0 cm. globulose, 8-9 mm. in diameter; seed coat latis, suborbicularibus, ellipticis vel obovatis, thin, membranous, closely adhc;ring to the apice obtuso vel apiculato rare retuso, basi peri carp and free from the surface of the coty­ subcuneato, laminis coriaceis concavis, mar­ ledons; cotyledons either completely fused gine integris valde revolutis, supra olivaceis or only partly so, or entirely free. minute pustulatis novellis dense subadpressi­ Type: Oahu, "Northern slope of Kaipapau pilosulis, in aetate nitidi-glabrescentibus, sub­ Valley," Degener and Park 4,169 (BISH). tus pallide flavidi-viridibus minute glandu­ DISTRIBUTION: Endemic to the island of losi -punctatis dense subadpressi -puberulenti­ Oahu. bus, petiolis 1-3 mm. longis 1 mm. latis Specimens examined: dense brunneis subadpressi-pilosulis, floribus OAHU: Koolau Mts.: Pupukea, dry gulch solitariis in bracteorum axillibus, bracteis side, Sept. 28, 1925, Judd 20; Pupukea, elev. subulatis dense brunneis subadpressi-pilosulis 300 m., Nov. 2,1925, Brown 1, 274; Kahuku, 1-1.5 mm. longis 1 mm.latis, pedunculis 5-7 entrance ofPupukea-Kahuku trail, lower edge mm. longis 0.5 mm. latis subadpressi-pilosu­ of decadent dry forest among lantana, Nov. lis, calycis tuba 2-3 mm. longa 3-4 mm. 22, 1931, Degener, Park, and Kwon 7,297 diametro subadpressi-pilosula, a 2 bracteis (BISH, NY); Laie, Kahawainui Gulch, elev. persistentibus subulatis brunneis subadpressi­ 100 m., March 2, 1928, Judd 71; Hauula, pilosulis subtensa, bracteis 1.5-2.0 mm. Papali Gulch on trail, March 1933, Judd; longis, calycis lobis 4 ovatis viridibus carnosis Hauula, on top of small cliff, elev. 160 m., persistentibus longitudine inaequalibus, quo­ Sept. 8, 1926, Judd 54; Kaipapau, northern rum uno jugo opposito 2 mm. longo 2 mm. slope of Kaipapau Valley, moderately dry lato, altero 3 mm. longo 2 mm. lato, petalis woods near top of ridge, Oct. 11, 1931, 4 albis in disci annularis margine insertis Degener and Park 4,169; Waimea, N. fork of ovatis vel obovatis 4 mm. longis 3 mm. latis Kamananui stream, very steep, north facing membranaceis ciliatis, apice obtuso, basi valley wall, alt. 750 ft., April 16, 1949, St. John truncato vix puberulo minute glandulosi­ 23,683, and Sept. 5, 1952, Wi/son, St. John and punctato, pistillo tereti subulato 2-5 mm. St. John 142, and Nov. 18, 1952, Wi/son and longo basi dense puberulenti supra glabroso, Lamberton 163; gully having prominent dyke, stigmati peltato, fructibus ovoideis sym­ north-north-east of Puu Kamaohanui, Dec. metricibus in sicco 8 mm. diametro dense 11, 1932, Degener 7,302 (NY). adpressi-pilosulis rubris ins igniter flavidi­ Closely related to Eugenia rariflora, E. koo­ glandulari-punctatis calycis lobis persisten­ lauensis may be clearly differentiated by its tibus coronatis. 176 PACIFIC SCIENCE, Vol. XI, April, 1957

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FIG. 1. Eugenia Inolokaiana. a, Habit X .50; b, flower bearing branchlet X 2; c, bud X 4; d, dissected flower bud X 8. From Rock 17,144B. Eugenia in Hawaii - WILSON 177

Tree 2.5-3 m. high; branches greyish­ above; stigma peltate; fruit ovoid, symme­ brown, glabrate, in age the bark smooth trical, 8 mm. in diameter (when dry), densely yellowish-red; leaf scars 1-1.5 mm. wide, appressed-pilosulous, red, conspicously yel­ rounded shield-shaped, reddish-brown; leafy low glandular-punctate, crowned by the per­ branchlets 0.5-1.5 mm. in diameter distinctly sistent lobes of the calyx. 4-angled to terete, densely brown subap­ Type: Hawaiian Islands, Molokai: Mauna­ pressed-pilosulous; internodes 7-24 mm. loa; June 1918, J. F. Rock 17,144, in the long; leaves 2.0-3.0 cm. long, 1.4-2.0 cm. Bernice P. Bishop Museum Herbarium. wide, suborbicular, elliptic or obovate; apex Common name: "Nioi." obtuse or apiculate, rarely retuse, base sub­ DISTRIBUTION: Known only from the type cuneate; blade coriaceous, concave, margin locality, Maunaloa, Molokai, and now prob­ entire, strongly revolute, above olive-green, ably extinct. minutely pustulate, densely subappressed­ Specimens examined: pilosulous when young, becoming glabrous MOLOKAI: Maunaloa: April 1918, Rock and shiny in age, below pale yellowish-green, 17,144; June 1918, Rock 17,144; Feb. 1920, minutely glandular-punctate, densely sub­ Rock 17,144B (BISH, GH). appressed-puberulent; midrib shallowly im­ Eugenia molokaiana is known only from the pressed above, elevated below, primary lateral collections made by Rock at Maunaloa, Mo­ veins alternate or opposite, 5-7 on a side, 4-9 lokai. When the locality was first visited in mm. apart, irregularly ascending at 140-150°, 1918, Rock photographed the tree which was meeting in an irregularly lobed intramarginal then already dying (Fig. 2). Since 1920 there vein 1.0-1.5 mm. from leaf margin, raised on has not been a single subsequent collection both surfaces but more distinct below, the of this species. In 1953 Rock was fortunate in veinlets less conspicuous, raised-reticulate; being able to revisit Maunaloa, Molokai, in petioles 1-3 mm. long, 1 mm. wide, densely search of this tree. Unfortunately there is no brown subappressed-pilosulous; flowers sin­ longer any trace of its existence. E. molokaiana gle, in the axils of bracts; the bracts subulate, has disappeared, like many other trees from densely brown subappressed-pilosulous, 1-1.5 that locality. mm. long, 1 mm. wide, peduncles 5-7 mm. If ever it should be found, E. molokaiana long, 0.5 mm. wide, subappressed-pilosulous; may be easily recognized by its small red calyx tube 2-3 mm. long, 3-4 mm. across, fruits, its peltate stigma, and its small, con­ subappressed-pilosulous, subtended by two cave, pubescent leaves. persistent subulate, brown, subappressed­ The description of the flower is based on a pilosulous bracts; the bracts 1.5-2.0 mm. bud just before anthesis. Unfortunately no long; calyx lobes 4, ovate, green, fleshy, per­ open flowers have been seen. The nature of sistent, of unequal length, 1 opposite pair the embryo and seed are not known since it 2 mm. long, 2 mm. wide, the other pair 3 mm. did not seem advisable to dissect the single long, 2 mm. wide; petals 4, white, inserted on fruit which was available. margin of annular disk, ovate or obovate, 4 Miss Marie C. Neal informs us that the mm. long, 3 mm. wide, membranaceous, nioi growing on Maunaloa, Molokai, was one ciliate, apex obtuse, base truncate, sparsely of three trees of that region which played an puberulous, minutely glandular-punctate; sta­ important role in Hawaiian traditions. Ac­ mens numerous (about 150); filaments white, cording to the notes in the Bishop Museum 0.5-2.0 mm. long, slender, subulate, glab­ on the native Hawaiian names of , the rous; anthers white, orbicular-ovate, 0.2-0.5 nioi from Maunaloa, Molokai, was identified mm. long; style terete, subulate, 2.5 mm. as "the tree form of a god, Kane-ikaulana long, densely puberulent at base, glabrous ula." The tree was used for making images 178 PACIFIC SCIENCE, Vol. XI, April, 1957

FIG. 2. Eugenia molokaiana growing on Maunaloa, Molokai. (Photo by]. F. Rock.) Eugenia in Hawaii - WILSON 179 by the command of the chiefs, or parts of it by Mr. Barclay. Hooker's Lond. Jour. Bot. were used in "vicious sorcery." The nioi 2: 211-240. growing on Maunaloa was also claimed to be DADSWELL, H. E., and H. S. INGLE. 1947. poisonous (Neal, letter). If our identification The Wood Anatomy of the Myrtaceae, 1. of this nioi as Eugenia molokaiana is correct, Trop. Woods 90: 1-7. then we can attribute the belief of its toxicity DE CANDOLLE, A. P. 1828. Prodromus system­ only to superstition. atis naturalis regni vegetabilis. Vol. 3: 1-494. Paris. TAXA REQUIRING FURTHER STUDY: DEGENER, OTTO. 1932-1934. Flora Hawaii­ ensis, fam. 273; Eugenia waianensis, July 15, Episyzygium oahuense Suesseng. and Ludw., 1932; Eugenia koolauensis, Aug. 10, 1932; Bot. Staatsaml. Mlinchen, Mitt. 1: 10,1950. Jambosa malaccensis, June 14, 1933; Jambosa The genus Episyzygium is based on a single Jambos, April 20, 1934. specimen which was collected by A. Meebold DEGENER, OTTO, and A. LUDWIG. 1952. in the Waianae mountains in 1930, [A. Mee­ Syzygium oahuense sp. nov. Bot. Staatsaml. bold 8,445 (M)]. I believe that the peculiar Miinchen, Mitt. 4:113. 4-loculed ovary is most likely an abnormality GAERTNER, JOSEPH. 1788. De Fructibus et of Eugenia sandwicensis. Until such a time that Seminibus Plantarum. Vol. 1: 182 + 384. additional corroborative material is collected, 79 pls. Stuttgart. I feel that the genus Episyzygium and the spe­ GRAY, ASA. 1854. Botany, Phanerogamia. cies Episyzygium oahuense should be considered Vol. 1 United States Exploring Expedition dur­ as being based on an aberrant individual. ing the years 1838 ... Charles Wilkes, U.S.N. Hillebrand (1888), Rock (1913), and Deg­ 15: 1-177. Off. ed. C. Sherman, Phila­ ener (1932-34) all reported a rare white form delphia. of Eugenia malaccensis ( Jambosa malaccensis f. HENDERSON, M. R. 1949. The Genus Eugenia cericarpa Deg.). I have not been able to locate (Myrtaceae) in Malaya. Gard. Bu!. Singa­ any specimens of this form. pore 12(1): 1-293. HILLEBRAND, WILLIAM. 1888. Flora of the Ha­ REFERENCES waiian Islands. 96 + 673, frontispiece, 4 AIRY SHAW, H. K. 1949. Additions to the maps. C. Winter, Heidelberg. flora of Borneo and other Malay Islands. INGLE, H. G., and H. E. DADSWELL. 1953. XX. Kew Roy. Bot. Gard. Bu!. Misc. Inform. The Anatomy of the Timbers of the South­ 1: 117-125. West Pacific Area. III. Myrtaceae. Austral. ALSTON, A. H. G. 1931. In Trimen, H. A Jour. Bot. 1(3): 353-401, 10 pls. Handbook to the Flora of Ceylon. Part 6 LAMARCK, ]. B. A. P. M. DE. 1789. Encyclo­ (supplement). vi + 350 pp. Dulau & Co., pedia methodique. Botanique. Vol. 3: viii + London. 759. Paris. AMSHOFF, G.]. H. 1942. Notes on the Myr­ LAN]OUW,]., and F. A. STAFLEU. 1954. Index taceae of Surinam. Bot. Mus. Univ. Utrecht, Herbariorum Part 1. 2nd ed. Regnum Vege­ Medde!. 86: 147-165. 2: 1-179. BACKER, C. A. 1945. Notes on the Flora of tabile Java. Blumea 5(3): 490-524. LINNAEUS, C. 1753. SPecies Plantarum. Ed. 1. BENTHAM, G. 1843. Enumeration of plants 1200 pp. Stockholm. collected by R. B. Hinds, Esq., and by Mr. MERRILL, E. D., and 1. M. PERRY. 1938a. Barclay in the Feejee Islands, Tanna, New On the Indo-Chinese Species of Syzygium Ireland, and New Guinea; to which are Gaertner. Arnold Arboretum, Jour. 19(2): added a few species gathered in Amboyna 99-116. 180 PACIFIC SCIENCE, Vol. XI, April, 1957

--- 1938b. The Myrtaceae of China. Ar­ --- 1913b. List of Hawaiian Names of nold Arboretum, Jour. 19(3): 191-247. Plants. Ter. of Hawaii, Bd. Agr. and For. --- 1939. The Myrtaceous Genus Syzy­ Bot. Bul. 2: 1-20. gium Gaertner in Borneo. Amer. Acad. A.rts SKEELS, H. C. 1912. In: Seeds and Plants and Sci., Mem. 18(3): 135-202. Imported during the Period from July 1 to MILLSPAUGH, C. F. 1900. Plantae Utowanae. September 30, 1911. U. S. Dept. Agr. Bttr. Field Columbo Mus., Bot. 2(1): 1-110. Indus. Bul. 248: 25. NIEDENZU, FRANZ. 1893. Myrtaceae. In SUESSENGUTH, K., and A. LUDWIG. 1950. Engler, A. and K. Prand. Die Naturlichen . Myrtaceae. Bot. Staatsaml. Munchen, Mitt. Pflanzenfamilien 3 (7) : 78-86. W. Engelmann, Leipzig. 1: 10. ROCK,]. F. 1913a. The Indigenous Trees of the WHITE, C. T. 1945. Contributions to the Hawaiian Islands. iv + 518 pp. 215 pIs. Queensland Flora NO.9. Roy. Soc. Queens­ Honolulu. land, Proc. 57: 21-36.