The Polynesian Species of Myoporum1

GRADY 1. WEBSTER2

INTRODUCTION trade, but neither then nor since has the naio THIS REVISION was originally begun as an attained any real commercial value. Brown analysis of the notoriously polymorphic (1935:. 279) reports that on the island ofRapa species Myoporum sandwicense of the Hawaiian (in southern Polynesia) naio wood is used Islands. After a considerable amount of work for building canoes and houses . had been done, however, it was found that The real value ofMyoporum to the Hawaiian the forms of Myoporum from southern Poly­ Islands resides in its role in the formation of nesia were very similar. Because of the close­ a dry forest cover and in the consequent ness of relationship, it seemed both more checking of soil erosion. On most of the is­ practical and more worth while to treat all of lands the dry forest region has been partially the Polynesian species together. or completely denuded, with serious conse­ In the Hawaiian Islands, Myoporum sandwi­ quent erosion. Reforestation work has been cense is an evergreen shrub or tree which carried out chiefly with quick-growing exotic grows mostly in dry forests on leeward slopes trees such as Casuarinaand Eucalyptus, but the from sea level to an altitude of 10,000 feet. appearance of the resulting vegetation is dis­ Its plasticity in habit is remarkable, as it may appointing from an esthetic point of view. It become a tree 15 meters high in the dry is to be hoped that eventually the original dry forests (Rock, 1913: 427) or a creeping , forest trees, such as native species of Dio­ fleshy, prostrate shrub on the beaches and low spyros, Acacia, Sophora, and Myoporum, will rocky islets. More rarely it may even invade regain a part of their lost dominance. Egler the rain-forest, as in Waikolu Valley on (1947: 425), studying the communities of the Molokai. dry southeastern section ofthe Koolau Range The Hawaiians used the species, which they on Oahu, predicted that although Myoporum called naio, for timber in building houses was rare at that time it might in the future as­ (Brigham, 1908: 83). Apparently it was not sume an important place in the Prosopis com­ cultivated, as it is in Rarotonga (Wilder, munity. 1931: 100), for the perfume of the flowers. Acknowledgments: Dr. Harold St.John ofthe There are references (Bennett, 1832: 257; University ofHawaii originally suggested this Hooker and Arnott, 1841: 93) concerning the study, made it possible to collect and observe attempts of traders of the 1820's to substitute the plants in the field, and has been of great naio wood for sandalwood in the Chinese assistance in criticizing and offering sugges­ tions for the manuscript. Mr. Otto Degener 'Part of a th esis submitted in partial fulfilment of re­ loaned his large and important collections for quirements for th e degree of Master of Arts in Botany at the University of Texas. Manus cript received Sep­ study and discussed with me .some of the tember 22, 1949. forms which I was unable to s~e in the field. 2Former grad uate assistant in Botany, University of Dr. B. C. Tharp readily consented to having Hawaii , and Teach ing Assistant in Biology, Unive rsity of Texas; now at the Herbarium, University Museum, the study continue at the University ofTexas, University of M ichig an. and he and Dr. W. Gordon Whaley kindly

[52 } Polynesian Species of Myopomm- WEBsTER 53

read and criticized the manuscript. I also wish that so far no really satisfactory subdivision to thank Miss M arie Neal for making avail­ of the genus has been made. able the facilities of the Bishop Museum The H awaiian species M . sandwicense was Herbarium, Miss Mary Lou Jeffrey and Mr. apparently first collected by Archibald Men­ William 1. Brudon for drawing the illustra ­ zies on Vancouver's expedition and was at tions , and the curato rs of the herbaria listed first referred 'to the New Caledo nian M . below for the loan of specimens : tenuifolium by H ooker and Arnott (1841: 93). Bishop Museum, H on olulu (BISH ) ; Bois­ It was described as a new species, Poly­ ser Herbarium, Geneva (G-BOIS); Otto coelium sanduiicense, ' by D e Cand olle (1847 : Degener, personal herbarium (DE G); Deles­ 705) and was redescribed in the genus sert Herbarium, Geneva (G-DEL); Gray Myoporum by Asa Gray (1866: 52). No other Herbarium (GH ); Royal Botanic Garden s, name was officially prop osed for any of the Kew (K ); New Yo rk Botanical Garden (NY) ; .Hawaiian plants until Leveille (1912: 63) Laboratoire de Phanerogamie, Paris (P); Riks­ published M. Fauriei based on a Faurie museum, Stockholm (S); University ofTexas , specimen from the island of Hawaii. Kraenz­ Austin (T); Naturhistorisches Museum, Vien­ lin (1929: 21)reduced the species to a variety na (W). but app ears to have misapplied the name to specime ns of var. sandwicense. TAXONOMIC POSITION Hugh Cuming had collected Myoporum in The genus Myoporum consists of about 30 the Austral Islands (Tubuai) in 1828, but species scattered over a wide area including over a hundred years went by before any new M auritius, Australia, N ew Zealand, New species were described from southern Poly­ Guinea, China, Japan, Micronesia, and Poly­ nesia. Cheeseman (1903: 291) listed as nesia. It is characteri zed by its relatively Myoporum sp. a M angaian plant culti vated

actinomorphic corolla and its fleshy drupe on Rarotonga. Wilder (1931: 100) I referred with usually solitary ovules in the cells. Like the Rarotongan plant to M . sandwicense, but other genera in the M yoporaceae, Myoporum Skottsberg (1933: 165) described it as a new has axillary flowers. .stamens with confluent species-M. Wilderi-and discussed its prob­ anther cells, pendent anatrop ou s ovules, and able relation ships to most of th e other in­ gland -dotted, alternate leaves . The Verbe­ sular species. Two years later Brown (1935: naceae is probably the most closely related 277) published th ree species from the Austral family, but it is well distinguished by its Islands and Rapa . ovules which are not apically attached and Variation in the Hawaiian myoporums has by its opposite leaves which lack internal been discussed by Skottsberg in his paper on 'secretory tissue (Solereder, 1899: 711). M . Wilderi and by Degener (1930: 261), The Polynesian species of Myoporum be­ but neither has published any new names long to section Pentacoelium of Gra y (1866: altho ugh D egener illustrated a pubescent­ leaved form and anno unced 'his intention of 51), which has priority over the more ap­ describing it in his Flora Hawaiiensis. propriate name Eumyoporum of Bentham (1870: 2). Wettstein (1895: 360) disting uish­ MORPHOLOGICAL CRITERIA ed three sections which contained Pacific Because of the extreme polymorphism of insular species- Pentacoelittm, Polycoelium, and many Hawaiian populations of Myoporum it Eumyoporum- but there is no justificatio n for is difficult to find morph ological characters placing these insular species in more than which do not vary as much between indi­ one section . Kraenzlin's sections (1929: 15) viduals as between varieties. An accurate are even more poorly founded; so it seems concept of the range of variation-arrived at 54 PACIFIC SCIENCE, Vol. V, January, 1951

by observation of a great many specimens­ often to be a response to wounding caused is essential in defining the taxonomic charac­ by grazing animals, although in some cases ters of the various groups. the lower "sucker" shoots from an uninjured Growth habit, as has been indicated, is so plant may bear pubescent leaves. In fact, greatly modified by environment that it is even if the leaves on basal or wound shoots probably of no taxonomic significance. The are not pubescent, they are often serrate and stem is significant only in distinguishing var. larger than normal leaves. Furthermore, there stellatum from var, Degeneri, the former hav­ is evidence that at least in some groups of ing pubescent br~nchlets. Myoporttm sandwicense the "juvenile" leaves There are many leaf shapes but few of of a: young plant are of the abnormal serrate­ them are constant in anyone population. pubescent type, while those formed later are Serrate margins are found sporadically among entire and glabrous. Further investigations nearly all the Hawaiian varieties and must be . on this subject are much to be desired. used with extreme caution as a taxonomic The number of flowers in the axillary character. On the other hand, this serrature is clusters varies not only between plants but constant in the southern Polynesian forms even on the same branch and thus does not and may be used dependably to separate M . appear to be of any great taxonomic value. Stokesii from M. rapense. Conspicuous dif­ The same can be said of the flower pediceJs, ferences in leaf texture occur among the which vary, widely, but inconsistently, in Hawaiian plants, but in the main these appear length and thickness. to reflect environmental differences. The The number and length of the calyx lobes, relative conspicuousness of the nervation within certain ranges of variation, are sig­ depends in turn on leaf texture and is thus nificant, while their shape (on account of its obviously unreliable. plasticity) is usually less so. The size and The leaves of all the Polynesian species shape of the corolla have not been used for have subepidermal pellucid-punctate tissue, distinguishing varieties; although there are and the distribution and type of these pel­ readily apparent differences, they are over­ lucid spots appear to be good specific cri­ lapping and inconstant. The amount of teria. M. Stokesii and M. rapense are difficult to pubescence on the corolla throat is usually separate from the New Zealand M . laetam quite variable and mayor may not have except for the fact that the glandular spots significance. . . on the leaves of the latter are far more Theovary is generally uniform in shape and conspicuous. In M. laetam the pellucid spots constant in size, within anyone variety. It is - are visible by reflected light (on the upper glabrous and has a somewhat expanded, dif­ leaf surface) and give a distinct "polka dot" ferently colored basal region which seems to effect in transmitted light. The spots of the be nectariferous, At anthesis the flowers may two Polynesian species are smaller, less con­ be noticeably fragrant, although I have spicuous in transmitted light, and almost personally found the flowers of M . sand­ invisible in reflected light. toicense to be almost odorless. In marked A readily recognizable and often valuable contrast to the Hawaiian species in this leaf character in Hawaiian Myoporutn is the regard are M. Stokesii and M. rapense, whose presence of pubescence. Unfortunately, while flowers are credited in collectors' notes as pubescent leaves are constant features of ranging in scent from "strong sweet" to varieties stellatum and Degeneri the y may " fetid." appear sporadically in populations of other The style furnishes some of the best tax­ varieties which ordinarily have glabrous onomic criteria, especially in its length and leaves. These sporadic occurrences seem in the presence or absence of pubescence. PolynesianSpecies of Myoporum-WEBsTER 55

This last character is undoubtedly valuable contain only one or two cells with well­ in delimiting species, although it may oc­ developed seeds. casionally be inconstant (on the islands of Hawaii and Raivavae). The abrupt basal TAXONOMIC CATEGORIES curvature of the style observed in some of The species is here considered to be a the specimens from southern Polynesia is an population whose individuals have in com­ inconstant character and its importance is monan ensemble of fundamental morpho­ hard to assess on the basis of the small logical characters different from that present amount of material studied . in other species. It is reasonable to suppose The number of stamens is one of the that this morphological distinctiveness is classical key characters for separating M. due to various types of barriers (such as sandioicense from the other Polynesian species, geographical, physiological, and genetical ) but .this distinction breaks down when a which prevent a free genetic exchange with large number of Hawaiian specimens is ex­ other species. amined . The length and heigh t of insertion The categories subspecies and variety, as of the stamens wer<,; found to be quite here used, apply to groups which occupy a inconstant, and the degree of divergence of discrete region within the total area of the the anther cells is dependent on age and is species and which are characterized by a lack of no taxonomic value at all. of sharp morphological distinctness from The drupes of most of the Hawaiian other groups within the species. A sub­ . varieties differ from the other Polynesian species differs from a variety in its greater species in having a greenish-white exocarp. morphological differentiation, but does not Some 'plants on the island of Hawaii (var. necessarily occupy a greater area. Both are Fauriei), however, may have either whitish, considered to lack genetic isolation-that is, pink, or purplish drupes. The drupes of M . varieties of the same species would be ex­ Stokesii and M . rapense are pink or red and pected to interbreed freely when brought brick-red or purplish-red, respectively, while together-and no doubt often owe their those of .M . laetum are said to be purple. existence to geographical barriers. J udging from the variability of color in M. In recent years there has been considerable sandwicense vat. Fauriei, however, it would be discussion among taxonomists as to whether unwise to' consider differences in shade of . the geographically delimited gro ups within a color as especially meaningful. The presence species should be given the rank of sub­ of color in the drupes of var. Pauriei and species or of variety. However, in this work its absence in the other Hawaiian varieties it has seemed better to use both categories do seem to be significant, however. in order better to accommodate the great The bony endocarp is usually subglobose sub-specific range of differentiation. or top-shaped and often ridged at the top or The form, in the present intrepretation, is a along the sides. Its shape, although extremely group of plants which occur within a variety variable, is a distinguishing character in some and differ by very minor or inconstant char­ varieties and species. The number of cells in acters; they mayor may not be geographically the drupe is inconstant, but a knowledge localized. Specimens may also be referred to of the range of variation will assist in placing forms when they are inadequate to give an a specimen in the proper category. Counting accurate concept ofthe population. Inasmuch the number of cells is not always easy as as the forms listed in this study are not con­ some of the abortive ones may be quite small. veniently separated morphologically and to There is often a considerable degree of ster­ some extent are designations based on in­ ility, for many drupes will be found to complete knowledge, it seems unwise to 56 PACIFIC SCIENCE, Vol. V, January, 1951 dignify them with Latin names. For con­ areas such as the Lualualei region on Oahu, venience they are listed by number, each western Molokai, and the Kohala peninsula under the island where it occurs. of Hawaii, where probably it was once very abundant. DISTRIBUTION In the Austral Islands the destruction of So far species of Myoporum have not been the native forests has been so complete that discovered on any of the ' islands occurring a clear picture of the distribution of Myo­ in the 3,OOO-mile expanse of sea between porum there may never be obtained. St. John Hawaii and the Austral Islands . It is not tells me that he made a special effort to col­ surprising that they are missing from low coral lect Myopo rum on these islands during the atolls such as Palmyra and Fanning, but their Mangarevan Expedition of 1934 and that absence from the Society Islands, the Mar­ few plants of the genus could have been quesas, and Makatea is unexpected, as is their overlooked in the vestiges of the native for­ absence from other large Pacific islands such est. Judging from his findings, M. rapense as Samoa and Fiji. var. Skottsbergii and the form described by The disjunct distribution of M. sandwi­ Brown as M. rimatarense may well be extinct. cense-with two subspecies in the Hawaiian On the basis of our present knowledge, Islands and one in the Cook Islands (Man­ however, it seems significant that M . rapense gaia)-is not unique, however. For instance, is found on the three lower islands of the Isachne distichophylla has been found only on Austral group and that its derivative M . Rarotonga and in the Hawaiian Islands . Stokesii is restricted to Raivavae. I Hedyotis section Bikkiocarpa has two species, One of the chief difficulties in under­ one from West Mauiand one from Rapa (see standing the distributional pattern of Poly­ Fosberg, 1943: 25). The genus Phy//ostegia. nesian Myoporum is that the means of long­ has 23 species in the Hawaiian Islands and range dispersal of the genus are still a matter one in Tahiti. Our present state ofknowledge of conjecture. It does not appear likely that is insufficient to explain these facts of dis­ the fruits are spread by ocean currents , since tribution. However, the theory of Zimmer­ my own observations show that they will man (1948: 49- 52) that high volcanic islands, float for no more than a few days. Dispersal now reduced to reefs or atolls by erosion, by birds is a possibility, since some endemic once existed as stepping-stones in the mid­ Hawaiian birds were known to have eaten Pacific indicates how 'these cases of bicentric the seeds; but there is no evidence that any distribution might have arisen. migratory birds are fond of the fruits. It must At the time of Captain Cook's visit be admitted that the dispersal mechanism of Myoporum undoubtedly existed on all seven Myoporum still remains unknown, and that ofthe main Hawaiian Islands. It is now absent the accidental transport by hurricanes is as from the denuded island of Kahoolawe, but likely a possibility as any. Forbes (1913: 86) learned from old visitors to the island that the naio had been seen there RELATIONSHIPS OF THE SPECIES before the dry forest was wiped out. The At present, and in spite of a fairly recent relative abundance of the species at those monograph (Kraenzlin, 1929), the genus localities where it still persists has no doubt Myoporum is in great need of critical revision . undergone great changes. In some areas, as The seat of the greatest systematic difficulty at Kaena Point on Oahu and the Puuwaawaa is the complex of chiefly insular species which district of Hawaii, it may be as common as it includes the three Polynesian species. Almost was before the dry forests were disturbed; nowhere within this insular complex are the on the other hand it is rare or absent from species well defined, and it is very difficult to Polynesian Species of Myoporum-WEBsTER 57 find morphological characters sufficiently It is pretty clear that the group of M. constant to distin guish them . sandwicense showing the greatest resemblance Because ofits great range of morphological to the Austral Island species is ssp. Wilderi, variation, M. sandwicense poses a particularly although it is not possible to say whether or difficult problem in species definition. In its not the latter originated directly from or most typical form the species is well charac­ coordinately with the Austral Island species. terized by its flowers with five stamens and a Among the groups ofM. sandwicense from glabrous style, and by its whitish-colored the Hawaiian Islands proper, var. Fauriei drupes with five or more cells. To some ex­ shows the greatest resemblance to ssp. tent, however, everyone of these characters Wilderi. Because var. Fauriei is restricted to proves to be inconstant when the complete the island of Hawaii, it seems likely that this range ofvariation ofthe species is ascertained . island was the point of immigration of The other Polynesian species typically differ Myoporum . Degener (1930: 266), on the basis in having flowers with four stamens and of the occurrence of pubescent, serrate leaves ' pubescent styles as well as reddish-tinted on certain wounded plants from Molokai, drupes with fewer cells, but most of these has suggested that the direct ancestor of the characters are inconstant. However, M. sand­ Hawaiian forms had leaves of this type , How­ wicense not only approaches the other Poly­ ever, this phenomenon does not seem to be nesian species very closely, but also is hard a case of a reversion to an immediate ances­ to separate from M. tenuifolium, a species tral type, but rather the expression of a lying to the west and southwest in Micronesia potentiality which is widespread, inasmuch as and Melanesia. This latter species is charac­ independeni: occurrences of pubescent leaves terized by the small size of its flowers and are known from other Hawaiian islands as fruits; but there are forms of M. sandwicense well as from Australia, New Zealand , and whose reproductive organs are of about the New Caledonia . same dimensions. Ofcourse, in all these cases It seems evident that var. sandwicense, the species distinctions can be maintained by most widely distributed of the Hawaiian using rather complicated combinations of groups, is derived from var. Fauriei, while characters; for instance, M . tenuifolium can it in turn has given rise to varieties lanaiense, be separated from M. sandwicense by the fact Degeneri, and stellatum. The origin of ssp. that the forms of the latter which have small St.-johniiis more obscure. It could have come flowers and fruit do not have pink-tinted directly from the adjacent population of var. drupes . Such contrived species differences are Fauriei, but the rather great morphological admittedly unsatisfactory, but they will have differences between the two entities make to serve for the present . The temptation to this rather questionable. The history of combine various of the insular species should Myoporum on the island of Hawaii is made be resisted until the entire group is surveyed, even more mysterious by the fact that var. because only then will it be possible to define sandwicense is absent from this island, even the exact limits of M. sandwicense and its though it is presumed to have originated nearest congeners. there. This suggests that the original popu­ Because the classification of the insular lation of var. sandwicense on Hawaii has myoporums is still in a very imperfect state, become gradually modified into the group one hesitates to speculate as to the phylogeny now known as ssp. St.-johnii. This suggestion of the Polynesian groups. However, the fol­ is of course hypothetical, but it would explain lowing observations on the possible relation ­ the curious anomaly of the absence of var. ships of the Hawaiian groups seem worthy sandwicense from its presumed place of origin, of statement. as well as the occurrence on Hawaii of forms 58 PACIFIC SCIENCE, Vol. V, January, 1951

of ssp. St.Johnii very similar to var. sand­ flora of the Austral Islands, our knowledge of uucense. the gro ups there is rather incomplete. Never­ On each island of the Hawaiian group theless, it is clear that M . rapense is very independent modifications, some of which closely related to M . laetum of New Zealand are superficially similar and have resulted in a and is no dou bt derived from it or a similar sort of parallel evolution, have arisen in the type. On Rapa and Tub uai it has changed populations. This curious phenomenon , little from the original immigrant, but on which is characteristic of populations on Raivavae it has given rise to and been almost some other Pacific islands as well, has been supplanted by the polymorphic M . Stohesii, remarked o n by Degener (1930: 266), who which itselfis undergoing incipient speciation realized the independent origins of the hairy­ in the manner of M . sandwicense. leaved and "peach-leaved;' forms on the various Hawaiian islands. Even more per­ SYSTEMATIC TREATMENT plexing from the taxonomic point of view It is not pretended that the taxonomic ar­ is the occurrence of reversions of characters. rangement which follows is definitive. Fur­ Thus, in tracing the character of style ther collecting on some of the Hawaiian pubescence in the Polynesian species, we find Islands may necessitate a reappraisal of some that it is present in M . rapense, beginning to of the unnamed forms listed herein and disappear in M . Stokesii, absent in ssp. sho uld result in the discovery of additional Wilderi, but reappearing in ssp. St.-fobnii . forms and perhaps even new varieties. Ade­ Naturally this sort of behavior of morpho­ quate understanding of a highly polymorphic logical characters introduces great hazards in genus such as Myoporum requ ires accurate classifying and interpreting the phylogeny of field observations, and careful notes by col­ the Polynesian species. lectors concerning Rower color and scent, It is interesting that there seems to be fruit color, and leaf variation will be of great little evidence of inter-varietal hybridization assistance in future studies . in the Hawaiian Islands. No indubitably hybrid specimens have been seen except MYOPORUM Soland. ex Forst. f., Prodr. possibly from the island of Hawaii, and here 44. 1786. additional field work will be necessary to establish the actual hybrid nature of such Generic synonyms pertinent to the Poly­ specimens . The conclusion seems inevitable nesian species are: that evolution in Hawaiian Myoporum has Pentacoelium Sieb. & Zucc., Abh. Akad. been most strongly conditioned by geo­ Muench. 4(3): 151. 1846. graphical isolation, while inter-varietal hy­ Polycoeliam A. DC, Prodr. 11 : 705. 1847. bridization has not played an important part . Prinastrum Nutt . ex Gray, Am. Acad. Sci. The fast rate of differenriation and slow dis­ 6: 53. 1866 (as synon ym). persal rate of the populations have been Low or erect shrubs, rarely tall trees. Leaves acting together to produce an intense micro­ alternate (rarely opposite), sessile or with a evolution. The taxonomic difficulty of char­ winged petiole, linear to lanceolate or el­ acterizing M . sandioicense is a reflection of the liptic, entire or serrate, glabrous or rarely rapidity and recentness ofthis process. Within pubescent, often viscid when young. Flowers .this one species can be found groups at all axillary, solitary or clustered,ebracteolate­ stages of differentiation-from individual pedicellate; calyx usually deeply 5-lobed, variants to populations of almost specific green and herbaceous, gland-dotted. Corolla distinctness. usually 5-lobed, imbricate in the bud, sub ­ Owing to the ruination of much of the actinornorphic, campanulate to funnelform, PolynesianSpecies of Myoportlm-WEBsTER 59

glabrous without, glabrous or pubescent should be accompanied by normal flowering within, usually glandular-punctate. Stamens branches if the correct determination is to usually 4, rarely more, alternate with the be made. corolla lobes; filaments subulate or terete, A. Style glabro us, or if hirsurulous then 2-3.5 mm. glabrous or pubescent, adnate to the corolla; long and fruit not red -tinged ... 1. M . sandwicense anther sacs confluent. Ovary of various B. Style glabrous. .shapes, often conic-cylindric, usually 2-5 ­ C. Stamens usually 5 (less commonly 4 or 6); corolla glabro us or pubescent; leaves various, celled (rarely the cells more numerous); style not elliptic-spatulate and abruptly acute . simple, usually as long as or longer than-the ...... 2. ssp. sandwicense ovary; stigma convex, entire or obscurely D. Leaves glabrous. 2-5-grooved (rarely bifid); ovules solitary. or E. Endoca rp stron gly depressed, not or rarely paired in each cell of the ovary, scarcely ridged, the thick wall exceeding the cells in diameter. .. . 5. var. lanaiense anatropous, pendent from the apex of the cell. E. Endocarp of various shapes, often ridged, Fruit a fleshy drupe; endocarp bony, usually rarely stron gly depressed, and then the more or less subglobose but sometimes de­ wall thinner than the cells in diameter. pressed or laterally compressed; seeds spindle­ F. Calyx lobes mostly 1-3 mm. long ; corol­ la often pubescent within; style mostly shaped, with scanty endosperm, the radicle 1.5- 3 mm. lon g; drupe always greenish­ supenor. white ; end ocarp 2- 6 mm . long, 4- 10­ celled ...... 3. var. sandwicense TYPE SPECIES : Myoporum laetum Forst. f., F. Calyx lobes mostly 3- 5 mm. long; corol­ here chosen as lectotype for the genus. It la glabro us (or rarely sparsely pubescent) should be noted that Forster attributed the within; style mo stly 3-5 mm . long; authorship of the genus Myoporum to Solan­ drupe sometimes pink or pur plish ; endo- carp 5-10 mm . long, 4-6-celled . der alone, and not to Banks and Selander, . . .. . ; ...... 4. var. Faariei as subsequent writers have incorrectly done. . D. Leaves pubescent. G. H airs branch ed ; branchlers pubescent; Section PENTACOELIUM (Sieb. & ZUCc. ) A. leaves lanceolate or elliptic-lanceolate ...... 6. var. stellatum Gray, Proc. Am. Acad. Sci. 6: 51-52. 1866. G. Hairs un branched. Pentacoelium Sieb. & Zucc., Abh . Akad, H . Branchlets glabrous; leaves often broad­ ly elliptic, entire or remotely serrate .. Muench. 4(3) : 151. 1846 (as a genus )...... 7. var. Degeneri Polycoelium (A. DC.) A. Gray, op. cit. (as H . Branchlets usually pubescent; leaves a section ). Ianceolate. Eumyoporum Bentham, Flora Australiensis 1. Corolla not over 6 mm. long ; style less than 3 mm . long . 5: 2. 1870 (as a section) ...... var. sandwicense, pubescent form Insularia Kraenzlin, ' Fedde Repert. Sp. 1. Coroll a over 6 rnm, long ; style 3 mm . Nov., Beih. 54: 15-16. 1929 (as a section) . long or more ...... var. Fauriei, pubescent form Ovary mostly 2-4-celled; ovule, one in C. Stamens constantly 4; corolla densely pubes­ each cell. Drupe mostly subglobose, not cent; leaves elliptic-spatulate, abrubtly acute . . strongly laterally compressed ...... 9. ssp. Wilderi B. Style hirsutulous ; £lower parts mostl y 6- 8; end o­ carp usu ally F 12-celled, often sharply ridged . . KEY TO THE POLYNESIAN SPECIES . . :.: ...... 8. ssp. St.-]ohnii A. Style hirsutulous (rarely glabrate in M . Stokesii), This key is intended for use in determining 4-6 mm . long; stamens 4; fruit pink or red; endo­ specimens in flower and fruit. Sterile 'twigs carp 3-6-celled. from the base of a plant or from a wounded ]. Leaves entire; endocarp angled . . . 10. M . Stokesii branch may sometimes have serrate and/ or ]. Leaves serrate ; endocarp not or scarcely angled...... 11. M. rapense densely pubescent leaves very unlike the K . Calyx lobes not ciliate ...... 12. var. rapense norm al ones. Specimens of this unusual type K. Calyx lobes ciliate ...... 13. var. Skottsbergii 60 PACIFIC SCIENCE, Vol. V,' January, 1951

1. M yop orum sa ndwicense A. Gray, Proe. mm . long, 2-9 mm. broad, smooth or ribbed, Am. Acad. Sci. 6: 52-53. 1866. subglobose or variously flattened, hard and bony when mature. Seeds one in each cell, Polycoelium sandwicense A. D C., Prodr. 11: spindle-shaped, papery when dry, 2-2.5 mm . 706. 1847 (illegitimate name). long. Usually much-branched shrubs 1-3 m. The name Myoporum sandwicense dates high, but stature varying from prostrate . from its use by Asa Gray in 1866, the origi ­ (littoral form ) to trees 15 m. tall. Stem usually nal combination Polycoelium sandwicense A. glabrous with smooth green or olive-brown DC. being illegitimate under the International bark when young, gray and cracking into Rules of Botanical Nomenclature, Article chunks when older. Leaves alternate, often 60 (1). The generic name Polycoelium was lanceolate but varying from linear-lanceolate a substitution for the earlier Pentacoelium of to ovate, acute to long-acuminate, entire or \ Siebold and Zuccarini, and hence was super­ often crenate or serrate, decurrent-petiolate, fluous when published. According to Article 3.5-22 em. long, 0.5-4 em. wide; usually 69 ofthe rules, Gray's adoption of the epithet som ewhat fleshy but varying from submem- sandwicense is treated as a new name and not branous to stiff-coriaceous when dry; usually as a new combination. glabrous or more rarely pubescent. Flowers As here construed M . sandwicense em­ axillary, in clusters of 2-10, often 1 or more braces three rather diverse subspecies, each abortive; pedicels terete or flattened, 0.5- of which could well be treated as a species if 1.7 em: long. Calyx lobes 4- 9, mostly 5, their differences were constant. I have been 1-6 mm . long, broadly ovate to oblong reluctant to reduce M. Wilderito a subspecies, or lanceolate, green and herbaceous or but if this were not done consistency would sligh tly scarious on the minutely glandular- demand the elevation of ssp. St.Johnii to denticulate margins, glandular-punctate, usu- specific rank, and this action does not seem ally glabrous. Corolla 4-9- (mostly 5-) lobed, warranted . 4 .5-12 mm .long, campanulate to funnel form, usually glandular-punctate, entirely glabrous 2. My oporum sandwicense ssp. sand­ to densely pubescent within; color pure wicense. . white, white with pink-purplish splotches, or entirely pink; corolla lobes ovate-deltoid Myoporum sandwicense A. Gray, Proe. Am. to oblong, emarginate, obtuse, or apiculate, Acad. Sci. 6: 52-53. 1866 (in part, excluding 2-7 mm. long. Stamens inserted on the reference to Douglas collection). corolla , the same number as and alternate Prostrate shrubs to tall trees. Leaves mos tly with the corolla lobes or occasio nally 1 or 2 lanceolate, less commonly almost linear or fewer; filaments flatrened -subulate, 1.5-5 ovate or elliptic, never spatulate, 3.5-22 em. mm . long ; anthers 0.5-1.5 mm . long, the long, 0.5-4 em. broad, glabrous or pubesce nt. cells confluent by a common suture and thus Flowers in clusters of 2-10 (mostly 3-5). changing from parallel to widely divergent Calyx mostly 5-lobed, lobes broadly ovate to at anthesis. Ovary cylindric-conic to de­ oblong or lanceolate, 1-5 mm. long. Corolla pressed-conic, 1.5-4 mm. long, glabrous, mostly 5-lobed, glandular-punctate, glab­ green, 4-12-celled, the base with a yellow or rous or pubescent within, 4.5-12 mm. long. purple apparently nectariferous ring ; style Stamens isomerous with the corolla lobes, the terete or flattened, 1.5- 6 mm . long; stigma fifth one occasionally reduced or absent. convex, obscurely 2-5-grooved. Drupe fleshy, Ovary 4-10-celled, 1.5-4 mm . lorig; style the exocarp greenish-white or creamy-white, glabrous, 1.5-6 mm . long. Drupe creamy- or rarely pinkish or purplish; endocarp 2-10 greenish-white or rarely pinkish or purplish; Polynesian Species of M yoporum-WEBsTER 61

endo carp 2- 10 mm . long, 2-9 mm. broad, however, the following key is submitted with subglobose to depressed. the caution that a clear differentiation of the This extremely variable and complex sub­ forms is not possible. species is restricted to the Hawaiian Islands. Key to forms of var. sandwicense A. Leaves all serrate, glabro us; style mostly 3-4 mm. 3. Myoporum sandwicense ssp. sand­ long -...... M olokai Form 3 wicense var. sandwicense. A. Leaves ent ire or occasionally' serrate or pubesc ent PIs. I, 2-6; 11,16--1 7, 27- 30; III , 38-40 and sharply serrate; style usually nor over 3 mm. long. . Myoporum sandwicense A. Gray, Proc. Am. B. Leaves glabrous. Acad. Sci. 6: 52-5 3. 1866 (in part, excluding C. Sramens 4; style mostl y 2.5-4 mm . long ...... - _Oahu Form 5, Kau ai Form 4 reference to Douglas and Nuttall collections). C. Stamens mo stly 5, rarely more. Leaves mostly lanceolate to oblong or D . Style 2.5-4 mm. long; corolla glabrous, broadly campa nulate, the lobes 5-7 . . : . . . . ovate lanceolate, rarely sublinear or ovate, _. . .. __ ... . _...... Easr MauL Form 4-22 em. long, 0.5- 3.8 em. broad, glabrous D. Style mostly 1- 3 mm . long ; coro lla often (except in one form). Flowers in clusters of pubescent, th e lobes mo stly 5. E. Corolla lobes 6- 8, corolla pubesce nt .. . . 2-9 (mostly 3- 6) ; pedicels 0.3-1.5 em. long. . _ _. _ -. __ Oahu Form 4 Calyx mostly 5-10bed (rarely the lobes 4, 6, E. Corolla lobes mos tly 5, rarely 6. or 7); lobes broadly to narrowly ovate or F. Leaves large, 8- 22 ern, long, often ser­ oblong, 1- 3 (rarely 4) mm. long. Corolla rate; style 2- 3 mm. long ; corolla pubes­ cent...... Kauai Form 1 5-lobed (rarely the lobes 6-7), glandular­ F.Leaves smaller, 5-1 4 cm. long, mostly punctate, glabro us or pubescent within, 4.5- 8 entire; corolla glabrous or pubescent. ­ (rarely to 9.5) mm. long. Stamens usually 5, G. Endocarp strongly depressed. H . Calyx less than 2 mm . long . occasionally one reduced (more rarely the ...... O ahu Form 3 number 4 or 6-8). Ovary 4- 1O ~ cell e d but H . Calyx over 2 mm . long __.. _... . . mostly 5- 7-celled; style 1.5- 3 (rarely 4) - Wesr Maui Form 2 G. Endocarp more or less isodiametric, mm. long. Drupe creamy-white, never pink or at least not depressed. or purplish; endocarp 2- 6 (rarely 7) mm. 1. Leaves linear-lanceolate, 8- 10 times long, mostly subglobose or top-shaped, as long as broad; calyx 2 - 3 mm . long ...... M olokai Form 2 usually ribbed. 1. Leaves mos tly less than 8 times as LECTO TYPE: Isles Sandwich, Oahu, Remy long as broad. 462 in the Gray Herbarium. ] . Leaves broadly elliptic. K. Leaves rather membranous . . . This extremely polymorphic variety is ...... Kauai Form 3 found at present on all the main islands K. Leaves fleshy, learhery when except Hawaii and Kahoolawe. - It is the dry _. Oahu Form 1 J. Leaves mos tly lanceolare . hetero geneous residuum which remains after 1. Leaves long-attenuate _ . the more well-marked populations have been ...... Ka uai Form 4 1. Leaves acute to acum inate. removed, and some of the forms enumerated M .Cells of drupe 4- 5; corolla here may warrant reappraisal when better mos tly about 5 mm . long known. . , . . . .. -. M olokai Form 1 It M. Cells of drupe mo stly 5-7, has seemed best to arrange the various less commonly 8 or mo re, forms in a geographic sequence, first for the rarely 4; coro lla ofren long- sake of convenience and also because forms er than 5 mm . . . . . Nii hau Form; Kauai from different islands which appear morpho­ Form 2,Oahu Form 2, logically similar are most likely related not Lanai Form, West Maui to each other but to adjacent forms on the Form 1 B. Leaves mo stly densely pubescent and serrate, same island. To give an idea of the corres­ sometimes glabrous entire leaves on the same pondence and relation ships of the forms, plant ______. M olokai Form 4 62 PACIFIC SCIENCE, Vol. V, January, 1951

2

13 15

2CM, Polynesian Species of Myoporum-WEBsTER 63

Specimens examined (DEG); Waimea, Kumuwela [Kumuweia] HAWAIIAN ISLANDS (without specific lo­ Ridge, bottom of gulch near stream, alt. cality) : 3,500 ft., St. John, Fosberg, & Oliveira 13852 Bennett (K); Gaudichaud 39 (G-DEL, P); (BISH); southeast side Kumuwela Ridge; 1839, Gaudichaud (G-DEL); Macrae (K); moist open forest, alt . 1,100 m., Fosberg 12650 Menzies (K); Wilkes Expedition (NY); "Ha­ (BISH); Kaholuamanu, Forbes319.K (BISH); waii," Wilkes Expedition (NY). same locality [Kaholuamano), Rock? 92 NIIHAU: (BISH), Rock 2478 (BISH, GH, NY), 5258 At foot of plateau, South East, Stokes (BISH), 5259 (GH). (BISH, NY); south corner of plateau, Stokes Form 3. Leaves elliptic, rather abruptly (BISH); Kalaalaau Vall~y, alt. 50 ft., St. contracted ro the tip. John 22747 (BISH); Mokouia Valley, alt. Without specific locality, Remy 463 (GH). 500 ft., St. John 23592 (BISH); Paniau, 1 Form 4. Like Form 2 but stamens 4. mile S. of, crest of windward cliff, alt . 1,200 Without specific locality, Wawra 2299 ft., St.John 23603 (BISH). (W) . KAUAI : Form 5. Leaves attenuate-acuminate, entire. Form 1. Leaves large, 8-22 em. long, tend­ Stamens 5. Drupe 6-7 mm. long. ing to be serrate. Stamens 5. Without locality, Brodie (BISH); ridge Waimea, seacoast, Mann & Brigham 585 west of the Hanapepe River, Heller 2452 (in part) (G-BOIS, GH, NY) ; Waimea, (BISH, G-BOIS, G-DEL, GH, NY, P). Polihale, flat back of beach, St. John et al. OAHU: 22956, 22957, 22961 (BISH, T). Form 1. Leaves fleshy, elliptic, up to 3 em. Form 2. Leaves mostly smaller and entire, wide (probably a mere ecological form). 5.5-14 em. long. Stamens 5. Without locality, Remy 461 (GH, P); Without locality, Menzies (GH) ; Wawra Mokuleia, near sea, Degener 9730 (DEG, (G-DEL); Wawra2072, 2407 (W); Oct. 1916, NY); Moku Manu 1. (islet off Mokapu Rock (BISH). Waimea, seacoast, Mann & Point), Munro (BISH). Brigham 585 (in part) (BISH, GH); Napali Form 2. Leaves lanceolate. Stamens 5 or Coast, at top of beach west of Kalalau rarely 4 or 6. Drupe subglobose, 5-:-8-celled . Valley, St. John et al. 23208 (BISH, T); (up to lO-celled in specimens from Koolau barren ridge, Waimea Drainage Basin, West Range). Styles mostly 1.5-2.0 mm. long. Side, Forbes 982.K (BISH); Waimea, Milolii Without locality: Beechey Expedition (G­ Ridge, on bare windswept headland, St.John DEL, K), Remy 462 (GH), Wilkes Expedi­ & Fosberg 13 711 (BISH) ; Waimea, Polihale, tion (GH). rocky lowland, alt. 30 ft., St.John et al. 22955 Waianae Range: Kaena Pt., Bryan (BISH) ; (BISH, T); Haeleele Valley, Degener 9736 Kaena Pt ., Kuaokala, Cowan 726 (BISH); (DEG); northwest side of Waimea Canyon, Kaena, in front of beach, alt. 15 ft., Cowan along ditch trail at 3,500 ft., Degener 20515 732, 737 (BISH); Kaena Pt ., near sea,

PLATE I Leaf outlines of the Polynesian species of Myoporum. EXPLANATION OF PLATE M . sandwicense var. stellatum: (1) from Webster 1243, Oahu. M . sandwicense var. sandwicense: (2) from Forbes 148.Mo, Molokai; (3) from Degener 9730, Oahu; (4) from Fosberg 13153, Oahu; (5) from Degener 12184, Molokai; (6) from Heller2452, Kauai. M . sandwicense var. Fauriei: (7) from Webster & Wilbur 1775, Hawaii; (8) from Rock 8335, Hawaii. M . sandwicense ssp. Wilderi: (9) from Wilder 781, Rarotonga. M . sandwicense var. Degeneri: (10) and (11) from Degener 12183, East Maui. M. Stokesii: (12) from St. John & Fosberg 15956, Raivavae; (13) from St. John & Fosberg 15964, Raivavae. M . rapense var. Skottsbergii: (14) from Cuming 1430, Tubuai. M . rapense var. rapense: (15) from Fosberg 11525, Rapa. 64 PACIFIC SCIENCE, Vol. V, January, 1951

HAWAIIAN ISLANDS ASSEMBLED

FIG. 1. D istribution of M . sandwicense var. sandwicense. Solid dots indica re exact locality, ope n dots approximate locality.(Base map, courtesy of Bernice P. Bishop Museum.)

Degener, Park, & Nitta ~73 1 (DEG) ; Kaena Fosberg 12350 (BISH) ; Mt. Kaala, forest at Pt., Forbes 1652.0 (BISH, NY); bluff above 2,500 ft., Degener, M ttrashige, & Kerr 19736 Kaena Pr., alt. 60 m., Fosberg 13153 (BISH) ; (transitional to Form 4) (DEG); Puu Kan e­ by railroad, Kaena Pt., alt. 15-20 ft., Neal hoa, southeast ridge of south peak, alt. 1,900 (BISH); Kaena Pt., Rttss (BISH), Topping ft., St.John 23291 (BISH) . (DEG) ; Nihoa Gulch, Mokuleia, alt. 150 ft., Koolau Range : Maunalua, Kealakipapa Hume 307 (BISH) ; same locality, alt. 400 'ft., Valley, alt. 5 m., Fosberg10919 (BISH, DEG); u; (DEG); Nihoa Gulch, Kaena, base of Maunalua, east botto m of Kalama Valley, cliffs, alt. 400- 500 ft., St.J ohn 11140 (BISH) ; alt. 15 m., V. O. Fosberg 93, 95 (BISH); near beach 1 mile east of Kaena Pt., Webster Maunalua, head of Manuwaii Valley, alt. 180 1077, 1079, 1080 (BISH, T), 1078 (BISH ); m., V. o. Fosberg 85 (BISH); arid rocky cliffs 2 miles east of Kaena Pt., alt. 250-350 region west of Makapuu Head, Degener, ft., Webster 1086, 1091 (BISH, T), 1090 Park, & Hirai 9728 (DEG, G-DEL, NY); (BISH ); at the top of the first gulch east rocky lowland 1 mile southwest of Makapuu of Kaena Pt., alt. 400 ft., IJ7ebster 1092.A Point, alt. 10 ft., Dunn (BISH) . (BISH, T), 1092.B (near to Form 3) (BISH); Form 3.Like Form 2 but drupe strongly cliffs 1 mile east of Kaena Pt., alt. 400 ft., depressed. Webster & Cowan 1097 (BISH, T); near road 1 mile east of Kaena Pt., Webster & KrattSs Waianae Range : cliffs 2 miles east of Kaena 1096 (BISH); Kewaula Valley, arid rocky Pt., alt. 250-350 fr., Webster 1081 (BISH, T); valley, Degener, Park, & Nitta 9732 (DEG, Kawaihapai Valley, alt. 400 ft., Suehiro NY); arid ravine near Kawaihapai, Degener (BISH) . 2197 (DEG, NY); steep brushy wall of Fo rm 4. Like Form 2 but flower parts 6-8. gulch, south side Makua Valley; alt. 500 m., Waianae Range: Kawaihapai, arid rocky Polynesian Species of Myoporum-WEBsTER 65

region, Degener, Park, & Topping 9729 (DEG, (DEG); Waikolu, steep valley wall below NY). Puu Kaeo, wet woods, alt. 3,000 ft., St. Form 5. Stamens invariably 4. Style 2-3 John et al. 23431 (BISH); same locality, alt. mm.long. 2,800 ft., St. John et al. 23466 (BISH); same Waianae Mountains, Hillebrand (K); Wai­ locality, alt. 2,000 fr., St. John et al. 23441 anae Range, Hillebrand (K); Kealia Trail, (BISH); Manawai - Kahanui ridge, moist Selling 3913 (transitional to Form 2) (S); forest, alt. 600 m., Fosberg 13392 (BISH). same locality, steep brushy slope, alt. 300 m., Form 4. Leaves serrate, densely pubescent, Fosberg 12858 (BISH); Puu Kaala, Waianae­ or glabrous and entire leaves on the same uka, alt. 650 m., Fosberg 12614 (BISH) ; plant. same locality, alt. 2,000 fr., St. John 12930 Puu Kolekole, Forbes 181.Mo (BISH); (BISH) , IJVebster &. St. John 1501 (BISH, T); east fork of Kawela Valley, three distinct near Kolekole Pass, Selling (Bog Survey) 3914 forms growing together on dry rocky slope, (BISH, S); Popouwela, Forbes 1593 (BISH, Degener 12181 .A (serrate-pubescent), 12181.C NY); Puu Kanehoa, east ridge, alt. 1,700 ft., (entire, pubescent), 12181 .D (entire,glab­ St. John 14065 (BISH) ; Lihue Gulch , east rous) (DEG). branch , Forbes 1688.0 (BISH) ; same locality, LANAI: Russ (BISH); Pouilihale ridge, semi-moist Without specific locality, Forbes (BISH); brushy ridge, alt. 700 m., Fosberg 13782 flats above head of Hawaiilanui Gulch, alt. (BISH); Palehua, Honouliuli, crest of ridge, 500 m., Fosberg 12523 (BISH); Maunalei alt. 2,000 ft., St.John 11162 (BISH); Palehua, Gulch , dry basalt talus, alt. 800 ft., St. John, alt. 650 m., Skottsberg 324 (S); near Mauna Eames, & Hosaka 18812 (BISH); Limestone . Kapu, Degener 9741 (DEG). Point, Munro (BISH) (identification doubt­ MOLOKAI: ful). Form 1. Leaves narrowly lanceolate, entire, : WEST MAUl: glabrous; drupe 4-5-celled. Form 1. Leaves narrowly lanceolate; drupe Kaunakakai, below government road, on subglobose or broadly ovoid. beach, Wilder (BISH, NY); east fork of Kawela Valley, Degener 12181.B (DEG); Without definite locality, Mann & Brigham Kamalo, alt. 1,000 m., Faurie 678 (BISH); 387 (G-DEL, GH, NY); Lahainaluna, Aug. Kamalo road, sea-level, Rock 12578 (BISH); 1910, Forbes (BISH, NY); south ridge of beach · at Kamalo, Rock 17137 (annotated Laniupoko Valley, alt. 1;000 fr., St. John & by Rock with an unpublished varietal name) Catto 17705, 17706 (BISH); Olowalu Valley, (BISH); Mapulo'u [Mapulehu], Rock 6153 Forbes 2325.M (BISH, NY); from Papawai (GH, NY); Pukoo, Faurie 679 (P); south Point toward Puu Anu through Manawainui slope of Halawa Valley, Degener 9737 (DEG, Gulch , Degener 9742 (DEG, NY); Pohakea G-DEL, NY). Gulch, arid windy slope, Degener9743 (DEG, G-DEL, NY); Wailuku, gulch southofWai­ Form 2. Leaves Iiriear-Ianceolate, entire, kapuu [Waikapu] Valley, Skottsberg 783 . glabrous; drupe 5-8-celled. (BISH, S). Slopes of Puu Kolekole, Forbes 148.Mo (BISH, NY, P). Form 2. Like Form 1 but drupe strongly de­ pressed. Form 3. Leaves oblong-lanceolate, all ser­ rate, glabrous; drupe 4-5-celled. Honokahau [Honokohau] Drainage Basin, Forbes 477.M (BISH, NY, P). North slope of Puu Kaeo, Degener 9739 (DEG); above head of Waikolu Valley, EAST MAUl: rainy shrubby gully, Degener 12182, 12184 The single form is much like West Maui 66 PACIFIC SCIENCE, Vol. V, January, 1951'

Form 1 but the style is longer and the flower The most perplexing forms of var. sand­ parts may be more than 5. wicense are those found on Molokai, and addi­ "Sandwich Islands," Hillebrand (probably tional field work must be done before a final from East Maui)(GH); East Maui, Hille­ systematic arrangement of them can be made. brand (K) ; Haleakala, Hillebrand (K); K ula, Molokai Form 3 is on the borderline of open rocky Kula land , alt. 1,000 ft., Wilder varietal status, but it 'is being kept as a form 282 (BISH) ; near Ulupal akua, Degener 9740 at present because its best character-con­ (DEG,G-DEL, NY); southwest slope of stantly and sharply serrate leaves- does not Haleakala, above Kanaio, Alexander & Kel­ distin guish it from populations on other logg 5353 (BISH) ; Hokukano, arid lava waste, islands within which serrate leaves are not Degener & Clay 19408 (DE G) ; , Nakaaha, uncommon . dryish lava waste, Degener 19306 (DEG); Nuu, rough lava flow, Forbes 191O.M (BISH) . , 4. Myoporum sandwicense ssp. sandwi­ The forms ofvar. sandwicense range in value cense var. Fauriei (Levl.) Kraenzlin, Fedde from those which are probably ecological Repert. Sp. Nov. Beih. 54: 21. 1929. modifications to thos e which approach varie­ PIs. I, 7-8; II, 18-19, 31; III, 41. tal distinction . Oahu Form 5 was at first Myoporrtm Fauriei Levl., Fedde Reperr. Sp. thought to be possibly a variety because ofits Nov. 11: 63. 1912. flowers with only four stamens, but the speci­ Leaves ovate -lanceolate to ovate or ellip­ men collected by St. John (number.23291) tic, acute or acuminate, entire or serrate, 5-17 from the center of its range at Puu Kanehoa cm. long, 0.9- 4 cm. broad, mostly coarse and shows that the stamen character is not con­ brittle-coriaceous but often thinner, glabrous stant. or rarely pubescent. Flowers 2-5 to an axil; HAW AII AN ISLAN DS ASSEMBLED

/

FIG. 2. D istribution of the other H awaiian groups of M. sandwicense. Circles represent var. Fauriei, squares ssp. St.:fohnii, erect trian gles var. stellatum , inverted triangles var. Degeneri, semicircles var. lanaiense. Solid symbols indi cate exact locality, open symb ols ind icate approximate locality; specime ns of really doubtful locality and th ose examin ed after the com pletion of th is plate are not mapped. (Base map, courtesy of Bernice P. Bisho p Museum.) Polynesian Species of M yoporum-WEBSTER ' 67 pedicels 0.5-1.7 em. long.Calyx lobes 5 "Hawaiian Islands"(no definite island (rarely 6), 2.5-6 (mostly 3-5) mm. long, cited): Collector unknown (BISH ); Bonite ovate -lanceolare to lanceolate. Corolla 5-lobed voyage, Gaudichaud (GH); Hillebrand (K ); .(rarely 6-lobed), 6.5- 12 mm.long, glabrous or Mann & Brigham 568 (BISH) . more rarely sparsely pubescent. Stamens Hawaii, without definite locality, Chal­ mos tly 5, rarely 6, often with one stamen re­ lenger Expedition (K), Wilkes Expedition (GH) ; duced or sometimes entirely wanting. Ovary South Hilo District, Waiakea, 22 miles up 2.5--4 mm. long, 4--6-celled (rarely 7-celled); Saddle Road, alt. 5,100 ft., St. j ohn, Cowan, style 2.5--6 (rnqsrly 3- 5) mm . long, glabrous. & Rogers 22391 (BISH); North Hilo District, Drupe creamy-white or often pinkish or Humuula, Puu Huluhulu, alt. 6,750 ft., Bryan rarely purplish; endocarp broadly ovoid, 5­ (BISH); same locality, alt. 6,760 fr., St. j ohn 10 mm . long, not prominently ribbed. et al. 23934, 23935 (glabrous sucker shoot TYPE: "Sandwich : Hawai ,M aunakea, from same plant as 23934), 23936 (pubescent 2,000 m ., juill. 1909 (Faurie, 677)." An iso­ sucker shoot from same plant) (BISH); Hu­ type is in the Bishop Museum. muula, Saddle Road 1 mile south of Omao­ This variety is limited to the upper dry koili, south slope of cinder cone, alt. 6,650 forest (altitude 4,000 feet to the timber line) ft., St. john, Cowan, & Rogers 22379, 22380 on the island of Hawaii." It is conspicuous (BISH); Pohakuloa, alt. 6,500 fr., Bryan because of its large leaves, flowers, and fruit (BISH) ; slopes of Mauna Kea above Waikii, and seems a distinct entity in spite ofits over­ Rock 8335 (BISH, GH); Puuwaawaa, Austin lap of characters with var. sandwicense. (BISH), Munro (BISH) ; Kona, above Hue­ Usually var. Fauriei has longer calyx lobes, hue, Rock 3489 (BISH , GH); H ualalai, Puu styles, and drupes than var. sandwicense, but Laalaau, alt. 6,300 ft., Kondo (BISH); Puu this does not always hold and then characters H ualalai, 4,100-6,500 ft. alt., St. john et al. such as the number of cells in the drupe, the 11394 (BISH); Kona, Captain Cook, south of lack of corolla pubescence, and the general Papaloa, alt. 5,000 fr., Degener & Murashige leaf aspect of var. Faurieimu st be considered. 20335 (DEG); Kealakekua, Greenwell Ranch, Var. Fauriei appears to be the most primitive alt. 4,500 ft., St. j ohn & Hatheway 23982 gro up of ssp. sandwicense and closely resem­ (BISH) ; South Kona District, Alika Home­ bles ssp. Wi/deri in flower size and drupe steads , Koa Mill, alt. 4,500 ft., St. j ohn et al. characters. Its retention of the pinkish drupe 22486 (BISH) ; Kilauea, Forbes ? (BISH), color and its flowers which ofte n lack the Forbes, Brigham, & Thompson (BISH, P); Ha­ fifth stamen are primitive characters ofspecial waii National Park, Kipuka Ki, alt. 4,300 ft., significance. Within var. Fauriei one finds a Webster & Wilbur 1702, 1703, 1704, 1705, great deal of polymorphism, there being large 1706, 1707 (BISH , T) ;Hawaii National Park, and small flowers, long elliptic-Ianceolate and Bird Park [Kipuka Puaulu], alt. 4,000 ft., short ovate leaves; but I have not been able Degener 9735 (DEG, NY); same locality, Bog to segregate these types into populations. Survey 3254 (S), Eastwood (NY) , St.john et al. Specimens examined 11260, 11266 (BISH), Setchell (S), Skottsberg HAWAII: 541 (BISH, S), Webster & Wilbur 1775, 1776, Form 1.Leaves glabrous; flower parts in 1777 (BISH , T). 5's; cells of drupe 4- 6. Form 2. Leaves pubescent; flower parts in 5's; cells of drupe 4- 6. 3Kraenzlin (1929: 21) erred in referring a collection from the New Hebrides (H erb. Boissier) to this variety . North Hilo District, Humuula, Puu H ulu­ I have seen the mate rial, and although it is too frag­ mentary to determine positively, there is no reason to hulu, open woo ds on cinder hill kipuka, alt. believe that it represents M . sandwicense. 6,570 fr., St.john et al. 23944 (BISH). 68 PACIFIC SCIENCE, Vol. V, January, .1951

Form 3. Leaves pubescent; flower parts 6--7; Specimensexamined cells of drupe 7-9. LAN AI: Puuwaawaa, Rock 3800 (BISH, NY). In Koa dry forest, Lanai, Munro 61 (BISH , several respects this form is intermediate be­ NY); plateau on lee side ofisland, alt. 600 m. tween vat. Fauriei and ssp. St.Johnii. (?), Wilder 90 (BISH ); xerophytic forest near Kanepuu, Fosberg 12551 (BISH); K anepuu, 5. M yoporum sandwicense ssp. sandwi­ K aa, alt. 1,600 ft., St. j ohn, Eames, & Hosaka cense vat. lanaiense Webster, vat. nov. 18803 (BISH) ; same locality, alt. 1,700 fr., St. j ohn & Cowan 22624 (BISH) ; flats above . PIs. II, 21, 33; III, 43 head of Hawaiilanu i Gulch, alt. 500 m., Fos­ Drupa depressi-conica; endocarpiurn non berg 12525 . (BISH); Poomai J =Paoma?], costatum, Munro (BISH); Mahana Valley, Munro 171Y2 Leaves lanceolate, acuminate, entire, gla­ (BISH); M ahana forehills, Rock 8119 (BISH , brous, 6-13.5 em. long, 0.8-1.8 em. wide. GH , W); mountains near Koele, Forbes 91.L Flowers 2- 4 per axil; pedicels 0.7-1 em. long. (BISH) . Calyx lobes 5, 2-3 mm . long, ovate-acumin­ 6. Myoporum sandwicense ssp. sandwi­ ate. Corolla 5-10beq, 5-8 mm. long, glabrous cense vat. stellatum Web ster, vat. nov. or pubescent, the tube 2.5-4 mm . long. Ovary PIs. I, 1; II, 20, 32; III, 42 about 2 .mm. long ; style 1.8- 3 mm. long. Drupe prob ably whitish in color; endocarp Prinastrum cauliflorum Nutt. ex Gray, Am. depressed-conic or hem ispheric in outline, Acad. Sci. 6: 52. 1866 (herbarium name pub­ lished in synonymy). not or scarcely ridged , 3-4.5 mm . high , 6--9 mm. broad, 5- 7-celled. Folia hirtella, ramuli hirtelli; pili ramosi. Branchlets and leaves pubescent with TYPE: Mahana forehills , Lanai, July 1910, branched hairs, somewhat glabrescent with Rock 8119, in the Bishop Museum Herbarium . age ; leaves elliptic-lanceolate, usually entire, Isotypes are in the Gray Herbarium and 4-9 em. long, 0.7- 2.2 em. broad. Calyx mostly Naturhistorisches Museum , Vienna. 5-10bed, pubescent, sepals 1.2- 2.5 mm. long. This variety, named for the island on which Corolla 5-10bed (rarely 6-10bed), 5-7 mm. it occu rs, is apparently now restricted to the long, glabrous or pubescent within , the dry forest near Kanepuu at the northwest end glandular dots rather inconspicuo us. Stamens of th e island . Ordinarily it is easily distin- mostly 4 or 5, if 5 then often one reduced. .guished from vat. sandwicense by its strongly Ovary 2-2.5 mm. long ; style 2-2.8 mm.long. depressed endocarp. Rarely, plants of vat. Drupe creamy-white; endocarp sub globose, sandwicense, such as Webster 1081 from Oahu ridged, 3- 5 mm. in diameter, 5- 7-celled. and Forbes 477.M from Maui, may have de­ T YPE: Oahu, Ewa coral plain near Barbers pressed end ocarps which appear superficially Point, alt. 10 ft., Webster 1243, in the Bishop similar, but these are easily distinguished in Museum Herbarium (isotype in the Univer­ cross section by the ir thinner walls (compare sity of Texas Herbarium ). Figs. 39 and 43). This distinctive variety is now restricted to . The status of vat. sandwicense on Lanai ·is the low plain of emergent coral at the south­ rather uncertain.Judging from the few collec­ western corner of Oahu . It is known from tions at hand, it seems to occupy a lower alti­ Kauai on the basis of a single collection by tudinal zone than var. lanaiense, but further Nuttall, but has not been collected on that collecting will be necessary before its distri­ island in the century since then. If Nuttall's butional relationship to var. lanaiense will be­ rather questionable locality was correct, the come clear. variety has evidently become extinct on PolynesianSpecies of M yOpOf Um-WEBSTER 69

Kauai . On Oahu it originally grew from the Branchlets glabrous. Leaves mostly ellip­ mouth of Pearl Harbor to Barbers Point, but tic, less commonly lanceolate, pubescent, the it has been exterminated over much of this hairs unbranched, entire or serrate, 4- 9 em. range by the encroachment of military es­ long, 1-2.5 em. broad. Flowers 5-6 per axil. tablishments. In the vicinity ofthe lighthouse Calyx lobes 5, glabrous or pubescent, 1.5­ it is still quite abundant, however, and will 3.5 mm. long. Corolla glabrous or pubescent continue to persist unless the vegetation is within, 5-7.5mm. long. Stamens 5, rarely 4 entirely removed . Ecologically, it must be or 6. Ovary 2-3 .5 mm. long; style 2-5 mm. highly specialized, as it has become adapted long. Drupe color unknown, presumably to a calcareous substrate quite different from white; endocarp 5-9 mm. long, 5-7-celled. the lateritic or basaltic soils on which the TYPE : East Maui, north mauka of Ulupala­ other varieties grow. kua, Degener 12183, in the Bishop Museum Although this variety superficially resem­ Herbarium. bles var. Degeneri in the pubescent leaves, it This variety occurs on the dry leeward has been independently derived, probably slopes of Haleakala, East Maui, from near from Oahu Form 5 of var. sandwicense. This Ulupalakua to the Kaupo Gap; a form from relationship is especially evident in the fact Molokai is referred here for convenience. It that var. stellatum shows the tendency toward is named for Mr . Otto Degener, long a stu­ the loss of the fifth stamen which is fully ex­ dent of the Hawaiian naio, who illustrated it pressed in the form from the Waianae Range. in his book, Plants ofHawaii N ational Park. The name of this variety alludes to the ap­ Specimens examined pearance of the branched hairs, a character MOLOKAI: unique in the genus as far as I can determine. Probably near Kaluaaha, Degener 9738 Specimens examined (DEG, NY). KAUAI: EAST MAUl: Atooi [=Kauai], Nuttall (K) . North mauka ofUlupalakua, Degener 12183 OAHU: (BISH, DEG); Auhi [=Auwahi], Ulupala­ Honouliuli, Barbers Point coral plain, alt. kua, Munro 385 (BISH); Kaapilopilo and Ka­ 10 ft., Cowan 787, 788 (BISH); same locality, pakahawai gulches, Forbes 1940.M(BISH); Degener & Park 9727 (DEG), Egler 37-268 Waiopaa [=Waiopai] Ranch , Forbes 1875.M (BISH), Hosaka 1359 (BISH), Pollock (F. (BISH); Haleakala, Kaupo Gap, Forbes Brown's) 1278 (BISH) , Rock 17038 (BISH, 1105.M (BISH) ; Haleakala, south of Kuiki labeled with an unpublished varietal name), along east side of Kaupo Gap, Degener Setchell (GH) , Webster 1243, 1244, 1250 17600 (NY). (BISH, T), 1245 (BISH) , 1249 (T); between Although the plant from Molokai is placed Barbers Point and Pearl Harbor, along beach, in var. Degeneri for convenience, it probably Degener & Park 9733 (DEG); Ewa, below has arisen independently of the Maui plants Magnetic Observatory toward beach, Egler & and may represent a different variety. How­ Hosaka 37-419 (BISH); Puuloa, 1909, Stokes ever, until the confusing forms from Molokai (BISH) ; east ·side of Pearl Harbor, Stokes are better understood, it seems unwise to de­ (BISH). scribe any new varieties from there. 7. Myoporum sandwicense ssp. sandwi­ 8. Myoporum sandwicense ssp. St.-Johnii cense var. Degeneri Webster, var. nov. Webster, ssp. nov. PI. I, 10-11 PIs. II, 22, 34-35 ; III, 44 Folia elliptica hirtella serrata; pili simplici; Petala et sepala plerumque 6-8; stylus ramuli glabri. hirtellus; drupa plerumque 7-12-locularis. 70 PACIFIC SCIENCE, Vol. V, January, 1951

16 18 19 - 20

21 22 23 24 25

27 29 30

32 33

31

I III I t 5MM 35 36 37 PolynesianSpeciesof Myopofum-WEBsTER 71

Shrubs to small trees, 2-8 m. high. Leaves with Form 2, which may be almost indis­ mostly oblong-lanceolate, entire, glabrous, tinguishable from some forms of var. sand­ 4-15 em. long, 0.5-3 em. broad. Flowers 3'--8 wicense. This is especially true of specimens per axil; pedicels 0.6-1.2 em. long. Flower from the vicinity of Punaluu in the Kau Dis ­ parts varying from 5-9 but mostly 6-8, the trict, but because they occur in the same area sepals equal to or less than the number of as plants of Form 1 and apparently are inter­ corolla lobes. Sepals 1-3 mm. long. Corolla mixed with them, it seems preferable to re­ 5'--8.5 mm. long, usually densely pubescent gard them as merely aberrant individuals of within, gladular spots absent or very few. ssp. St.Johnii, which for some reason (possi­ Stamens usually 5-9 (very rarely less than 5), bly they are remnants of an ancient popula­ the same number as or less than the number tion of var. sandwicense ?) are especially of corolla lobes. Ovary 2-3 mm . long; style common in this vicinity . hirsutulous, 1.7-3.5 (mostly 2-3) mm . long. This group is named for Dr. Harold St. Drupe creamy-white; endocarp usually John of the University of Hawaii, who sug­ strongly ridged, more or less laterally com­ gested the study of Myoporum and whose pressed, 3-8 mm . long, 3.5-8 mm . broad, 5­ assistance has greatly helped to overcome the 13-celled but typically 7-12 ~celled. disadvantages of completing this work at a TYPE: Hawai~ , Kaupulehu, 2~3 miles east distance from the Hawaiian Islands; of Huehue, alt. 2,000 ft., Webster & Wilbur Specimens examined 1835, in the Bishop Museum Herbarium (iso­ HAWAIIAN ISLANDS (without specific lo- type in the University of Texas Herbarium). cality) : This polymorphic subspecies is restricted Douglas21 (K) , Douglas (W). to the lower dry forests on the island of Ha­ HAWAII: waii. In 'its possession of. a hirsutulous style Form 1. Style hirsutulous and flower parts it is unique among the groups within M. mostly 6-8. Cells of endocarp 7-13. sandwicense, and it is further well marked by North Kona, Alexander & Kellogg 5367 its large number of perianth parts, reduction (BISH); vicinity of Huehue near main road, of glandular spots on the corolla, and many­ BogSurvey 3213 (BISH, S); between Puuwaa­ celled, sharply ribbed, endocarp. Morpho­ waa and Huehue, Degener 2210 (DEG) ; Puu ­ logically, the typical form (Form 1) isfully as waawaa, Degener 9726 (DEG, NY), Nishina distinct from the other varieties of M. sand­ (Degener's) 9744 (DEG, NY); 2.3 miles north­ wicense as many species of this genus are from . east of Huehue Ranch House, Kaupulehu, one another. But the typical form intergrades alt. 2,000 ft., St. John et al. 22525 (BISH) ;

PLATE II Reproductive stru ctures of the Polynesian species of Myoporum. EXPLANATION OF PLATE Mature pistils (FIGS. 16-26). M . sandwicense var. sandwicense: (16) from Webster 1097, Oahu; (17) from Heller 2452, Kauai. M. sanduricense var. Fauriei: (18) from Webster & Wilbur 1706, Hawaii; (19) from Webster & Wilbur 1702, Hawaii . M . sandu/icense var. stellatum: (20) from Degener 9727, Oahu. M . sandwicense var. lanaiense: (21) from Rock 8119, Lanai. M . sand­ uiicense ssp. St.-]ohnii: (22) from Webster & Wilbur 1835, Hawaii. M . sandwicense ssp. Wilderi: (23) from Wilder 781, Rarotonga. M . Stokesii: (24) from St. John & Fosberg 15947, Raivavae. M . rapense var. rapense: (25) from St. John & Fosberg 15280, Rapa. M . rapense vat. Skottsbergii: (26) from Cuming 1430, Tubuai. Side view of endocarps (FIGS. 27-3 7). M . sandwicense var. sandwicense: (27) from St.John et al. 23208, Kauai; (28) from Degener 9729, Oahu; (29) from Degener 9737, Molokai; (30) from Forbes477.M, West Maui. M . sandwicense var. Fauriei: (31) from Webster & Wilbur 1706, Hawai i. M . sanduilcense var. stellatum: (32) from Degener9727, Oahu . M . sandwicense vat. lanaiense: (33) from Rock 8119, Lanai. M. sandwicense ssp. St.-]ohnii: (34) from Webster & Wilbur 1835, Hawaii ; (35) from Degener 2211, Hawaii. M . Stokesii: (36) from Fosberg 11784, Raivavae. M . rapense var. rapense: (37) from St.John & Fosberg 15280, Rap a. 72 PACIFIC SCIENCE, Vol. V, January, 1951

40 41

42 43 44

45 46 .. :~ ::l22)3

-; ".- .

I •• 5MM PLATE III Reproductive structures of the Polynesian species of Myoporum. EXPLANATION OF PLATE Endocarps in cross section. M. sandwicense var. sandwicense: (38) from V . O. Fosberg 93, Oahu; (39) from Webster 1081, Oahu; (40) from Degener 9737, Molokai. M . sandwicense var. Fauriei: (41) from Webster & Wilbur 1704, Hawaii. M . sandwicense var. stellatum: (42) from Degener 9727, Oahu. M . sandwicense var. lanaiense: (43) from Fosberg 12525, Lanai. M. sandwicense ssp . St.Johnii: (44) from Webster & Wilbur 1835, Hawaii. M . Stokesii: (45) from St. John & Fosberg 15956, Raivavae, M. rapense var. rapense: (46) from Fosberg 11525, Rapa.

Puuwaawaa, 4 miles southwest of Puu Ana­ lehu, 2-3 miles east of Huehue, alt. 2,000 ft., hulu, alt. 2,000 ft., Webster & Wilbur 1874, Webster & Wilbur 1835 (BISH, T); Keahuolu, 1883, 1895 (BISH, T); Puuwaawaa, 3 miles ' alt. 400 ft., on aa in dry scrub, St.John &Lane west of Puuwaawaa Hill, alt. 3,000 ft., Web­ 23978 (BISH) ; Kapua ?, near the sea, Bryan ster & Wilbur 1852, 1857 (BISH, T); Kaupu- (BISH); Kapua, on kipuka south of Hana- Polynesian Speciesof M yoporum-WEBsTER 73

keaumoe triangulation point, near sea level, Shrubs or small trees . Leaves spatulate­ Bryan 761 (BISH) ; Manuka Mauka, alt. 750­ elliptic or obovate-elliptic, acute or apiculate, 1,750 ft., St. John et al. 11326 (BISH) ; Kau, glabrous, entire, 5-10.5 cm. long, 1.8- 3.3 ern. halfway between Kaalualu and Waiohinu, wide. Flowers 3-4 per axil; pedicels 0.8-1 cm. Degener9734 (DE G, NY); old aa flow, Hilea, long, flattened.Calyx 5-lobed, glandular­ Russ (BISH); Kau D istrict, Ninole, Wailua dotted, glabro us, the lobes mostly 1.5-2 mm . Ninole, basalt flow near shore, alt. 15 ft., St. long. Corolla white, 7.5-9 mm . long, John, Hatheway, & M orton 23950 (BISH) ; glandular-spotted, pubescent within, 5-lobed. Punaluu, Degener & Wiebke2207 (DEG, NY) ; Stamens 4 (sometimes with traces of a fifth on aa flow, Punaluu, alt. 500 fr., N eal (BISH) ; according to Skottsberg). Ovary globose­ Hawaii National Park , Bird Park, Degener pyriform, about 3 mm. long; style 3-4.5 mm. 9725 (DEG) ; Naaulu Forest, Kealakomo, alt. long, curved at the base, glabrous. Drupe 1,700 ft., Fagerlund & Mitchell 800 (BISH) . color unknown (probably purplish or red­ St.John etal. 23950 has the hirsutulous style dish); endocarp top-shaped, about 5-6 mm . typical for the subspecies but is remarkable in long, 4- 6-celled. having the stamen number highly variable TYPE: Rarotonga, Wilder 781, in the Bishop and reduced to 2-3 in the majority of flowers. M useum Herbarium. The locality of Degener 9725 should be re­ This subspecies is cultivated by the natives garded with suspicion because Bird Park has of Rarotonga, in the Cook Islands of south­ been visited many times by botanists,' and ern Polynesia, and has been found in the wild only var. Fauriei has beencollected there. state only on the nearby island of Mangaia. Form 2. Style very sparsely hirsutulous or It is fairly well distinguished from the other glabrous; flower parts mostly 5-6; cells of two subspecies by its leaf shape and by its endocarp 5-7. flowers with only 4 stamens. However, mos t Seventeen miles from Kohala toward Wai­ of the characters given by Skottsberg (1933: mea, Degener & Wiebke 2208 (DE G, NY); 155) to separate M . Wilderi from M. sand­ Puu Keekee, Degener, Greenwell, & Mura­ wicenseare not applicable when a large amount shige 20020 (DEG) ; 20 miles from Waimea of material of the latter species is examined. Hairy corolla lobes occur in most of the Ha­ toward Kona, Degener & Wiebke 22 11 (DEG, waiian varieties, and the density of the pube­ NY); Puuwaawaa, Forbes 49.H (BISH); Kau scence may be as great as in the Rarotongan Desert 25 miles west of Kilauea, Degener & Wiebke 2209 (G-DEL, NY); Kau District, plant. .The constant absence of the fifth stamen in the Cook Island population seems Pun aluu , St. J ohn et al. 11316 (BISH) ; same like a good distinction, but the form of var. locality, St. John, Cowan, & Rogers 22426 sandwicense from the Waianae Range of Oahu (BISH) . usually lacks the fifth stamen, and similarly Degener, Murashige, & Greenwell ' 20019 reduced flowers are not uncommon in var. (DEG), collected near Na Puukulua, is sterile stellatum and var. Fauriei. The length of the and has serrate-pubescent " juvenile" leaves, style does not distinguish Skottsberg's spe­ but since it was found near number 20020 it cies because the style in var. Fauriei reaches probably belongs here. 5 mm . and more in length. There remains the leaf shape to fall back upon, and although 9. Myoporum sandwicense ssp. Wilderi this is rather distinctive in ssp. Wilderi-the (Skottsberg) Webster, comb. nov . rounded apex of the elliptic leaf being rather PIs. I, 9; II,23 abruptly contracted into an apiculate tip - it Myoporum Wilderi Skottsberg, Acta Horti is variable even on the type specimen and does Gotob. 8: 165-166. 1933. not seem to be a character of specific value. 74 PACIFIC SCIENCE, Vol. V, January, 1951

Perh aps the best distinguishing character from the probable ancestor, M . /aetum, and it for ssp. Wi/deriis that its flowers have an odor seems likely that the ancestral form whic h so noticeable that the plant is cultivated for populated the Austral Islands and Rapa was perfume. In th is respect the subspecies re­ very close to M . rapense var. rapense. sembles the stro ngly scented Austral Island This species is m uch more variable th an M . plants and contras ts with the relatively odor­ rapense, and some of its forms diverge widely less H awaiian plants . Th e basal curvature of from the norm. Fosberg 11737 is remarkable for the style is another link between the Cook the petalloid appendages which are present on Island and Austral Island plants; bu t this the stamens, but as no other specimen from character is probably dependent on the length the island shows this, the plant had better be of the style and is of no great intrinsic signi­ regar ded as a minor variant. ficance. The inconstancy of style pubescence in M . Specimens examined Stok esii raises difficulties in separating it from COOK ISLANDS: M. sandwicense; the forms of M : Stokesii with Raroto nga, Wilder 781 (BISH,N Y); Man­ glabro us styles (fill be posi tively distinguished gaia, Nov. 1928, Graham (BISH) . from ssp. Wi/deri only by their narrower leaf shape. The approach of these forms to ssp . 10.M yo p orum Stokesi i F. Brown, Bishop lVilderi illustrates the difficulty of defining Mus. Bu!. 130: 277- 278, fig. 41. 1935. specific lines in the insular myoporums. Even PIs. I, 12; II, 24, 36; III, 45 though the distinctions between the Hawaiian and Austral Island species may thus break Shrubs or trees, 1-5 m . high. Leaves elliptic­ down, however, the species are in the main lanceolate, entire, glabrous, 3.5- 9 cm. long, distinct, and it wo uld be un wise to combine 0.5- 1.7 cm. broad, acute, apiculate, or them before all the insular myoporums have attenuate-acuminate, alate-petiolate. Flowers been carefully studied. 1- 3 (rarely 4) per axil, sometimes accompan­ ied by 1 or more small abortive flowers; pedi ­ Specimens examined cels 0.7-1.4 em , long. Calyx 5-lobed, RAIVAVAE: glandular-dotted; calyx lobes 2- 4 mm. long, Form 1. Style copiously hirsutulous, entire-margined .Corolla pure white or pink­ Near mountain ,to p, alt. 1,000 ft., Chapin spotted in the throat, rather heavily fragrant, 871 (NY) ; Pic Rouge, northwest side, alt. 9-12 mm. long, pubescent within. Stamens 4. 120 m. , St.J ohn & Fosberg 15947 (BISH) ; same Ovary 2.5- 3.5 mm. long ; style hirsutulous locality, alt. 150 m ., St.John & Fosberg 15956, (rarely almost or quite glabrous), curved at 15964 (BISH) ; Maunanui, ridge, alt. 900 ft., the base, 4-6 mm. long when mature. Drupe Stokes 11 (BISH); M atotea, alt. 78 m., Stokes 90 translucent pink to red ; endocarp turbinate, (BISH, type-:- as to sheet from Raivavae) ; ridged, 4-6 (mostly 5) mm. long, 3-6-celled " Raivavae," Whitney expedition 237 (N Y). but mostly 4-celled. . Form 2 . Style hirsutulous ; stamens with T YPE : Raiv avae, Matotea, alt. 78 m., April petalloid app end ages. 28, 1922, Stokes 90, in the Bishop Museum Herbarium . Vaiuru, N . E. slope, alt. 30 m., Fosberg This species, known from a single island, 11737 (BISH) . is closely related to M . rapense. Its differences Form 3. Style sparsely hirsutulou s to gla­ are not very profound but appear to be con­ brou s. stant; for instance, no leaves of M . Stokesii South side of M t. Turivao, rock ledge, alt. have been found to show the serrations ofM . 240 m., Fosberg 11784 (BISH) ; south side of rapense. M . Stokesii is evidently more distant M t. Araua, alt. 300 m., St.John 16201 (BISH ); Polynesian Species of Myoporum-WEBsTER 75

Pic Rouge, northwest side, alt. 140 m.,St. 12. Myoporum rapense var. rapense John & Fosberg 15957 (BISH); Mt. Taraia, PIs. I, 15; II, 25, 37; III , 46 south side, alt. 250 m., St. John & Kondo Myoporum rapense F. Brown, Bishop Mus. 15996 (BISH). Bu!. 130: 278-279, fig. 42. 1935. 11. Myoporum rapense F. Brown, Bishop Leaves lanceolate or obovate-lanceolate, Mus. Bu!. 130: 278-279, fig. 42. 1935. 4-10 em. long, 0.7-2 em. broad. Calyx lobes lanceolate, acuminate, 2-3 mm . long; mar­ Shrubs or trees, 1-10 m. high . Leaves Ian­ gins entire or minutely serrulate. Corolla 8-9 ceolate or somewhat obovate, sharply serrate, (rarely 11-12) mm. long. Ovary 2.5-3 mm. glabrous, 4-11.5 em. long, 0.7-2 em. broad, long; style 4-5 (rarely 6) mm. long. Endo ­ acute to acuminate, alate-petiolate. Flowers carp 5-8 mm. long, 3.5-5 .5 mm. broad, 1-4 (rarely 5) per axil, often 1 or more abor­ 3-6-celled. tive; pedicels 0,5'-1.5 em. long. Calyx lobes 5, TYPE: Rapa, alt. 170 m., Sept. 13, 1921, glandular-dotted, lanceolate and acuminate, Stokes 161, in the Bishop Museum Herbarium. 2-3.5 mm. long; margins entire, minutely Brown (1935:279) cited two collections from serrulate, or prominently ciliate. Corolla 5­ Rapa as types. I have chosen the first one as lobed (rarely 6-lobed), white, pink, or purple­ the lectotype, since the number of the other spotted in the ,throat, with a heavy sweet or (Stokes 90) is the same as the type of M. sometimes fetid odor, 8- 12 mm. long, pube­ Stokesii. scent within . Stamens 4, all well developed or one a rudiment. Ovary 2.5-3.5 mm. long; Specimens examined style hirsutulous, curved at the base, 4-6 mm. RAIVAVAE: long . Drupe brick- or purplish-red; endocarp This form has a corolla about 12 mm. long, turbinate or oblong-ellipsoid, not or scarcely style 6 mm. long. ridged, 5-8 mm. long, 3.5-5.5 mm. wide, Vaiuru, 1 km. east, top of beach, alt. 0.5 m., 3-6-celled (often one or more cells empty) . Fosberg11723 (BISH). This species occurs on Tubuai, Raivavae, RAPA: and Rapa and is more constant morpho­ This form has a corolla 8-9 mm. long, logically than is its probable derivative M . style 4-5 mm. long. Stokesii. It is separated from this latter species Anarua, alt. 50 ft., Stokes 161 (BISH, type) ; by characters which are very stable for the Anarua Valley, southeast ridge ofMt. Perahu, genus Myopo rum, and only one specimen is alt. 300 m., Fosberg 11521 (BISH); Matauri in any way intermediate. The form of M . Point, near lower edge of forest, alt. 3 m., rapense on Raivavae has typical leaves and Fosberg 11525 (BISH); Anarua Bay; top of fruit but flowers more nearly the size of M. beach, alt. 1 m., St.John 15707 (BISH); Area, Stokesii. This form does not seriously break alt. 145 m., St.John & Fosberg 15280 (BISH); down the boundaries between the two species, Toutore, west of Mt. Vaitau, alt. 240 ' m., however, and it seems best to maintain them St.John & Maireau 15409 (BISH); northeast as distinct. ridge of Mangaoa Peak, alt. 320 m., St.John While M . rapense is clearly separable from & Maireau 15381 (BISH) ; north slope of Mt. M. Stokesii, it is not nearly so easily dis­ Lekie, mixed woods, alt. 250 m., St. John & tinguished from the New Zealand M . laetum. Maireau 15624 (BISH); Karapo Rahi 1. For the present, however, its more incon­ [islet off Rapa], alt. 75 m., St.John & Maireau spicuously punctate leaves (fully discussed 15600 (BISH) ; Taunoa Ridge, alt. 400 ft., under Morphological Criteria earlier in this Stokes 90 (as to sheet from Rapa) (BISH); paper) may serve to separate it. Hiri, north ridge from Lekie, alt. 500 ft., 76 PACIFIC SCIENCE, Vol. V, January, 1951

Stokes 217 (BISH, NY); Kulukulu, Hiri, ridge, Myoporum rimatarense F. Brown, Bishop Mus. alt. 170 fr., Stokes 304 (BISH) . Bul. 130: 280, fig. 43. 1935. This species from Rimatara, AustralIslands, 13. Myoporum rapense var. Skottsbergii is known from such fragmentary material (the Webster, var. nov. flowers being unknown) that its exact dispo­ PIs. I, 14; II, 26 sition is impossible. Judging from the leaf Lobae calicis ciliatae, shape it might be a form of M. sandwicense Leaves elliptic-lanceolate, 5.5-8 em. long, ssp. Wi/deri, but this is only a guess . St.John 1.2-1.7 em. broad. Calyx lobes ovate, acu­ tells me that the vegetation on Rimatara is minate-tipped, sparsely long-ciliate on the badly despoiled and that the Myoporum there margins, about 2 mm. long. Corolla 10.5-11 may well be extinct. mm . long. Ovary 2.5 mm . long; style 4-4.5 Myoporum tenuifolium Forst f., Prodr. 44.1786. mm. long. Color of flowers and fruits un­ Nadeaud (1897: 113) credits this species to known; endocarp 5.5-6 mm. long, with 2 Tahiti (as a cultivated plant) and to Raivavae . fertile and 2 sterile cells in the one fruit cut The Raivavae plant is probably M. Stokesii. open. The cultivated Tahitian plant may also be M. TYPE : "Toubouia" [=Tubuai], Cuming Stokesii, but I have seen no specimens. M. 1430, in the Kew Herbarium. tenuifolium is one of the most widespread This variety is known only from the type species of the genus, occurring from New collection made by Cuming in 1828. One Caledonia into Micronesia, but it has not been might explain its not having been collected collected in Polynesia proper. there again by assuming that the locality given on the label is incorrect, but St: John REFERENCES (1940: 88) has studied Cuming's voyages and cites his collection numbers 1423-1433 as BENNETT, GEORGE. 1832. An account of the being made on Tubuai. A more likely ex­ sandal wood tree (Santalum), with observa­ planation is that the variety is extinct, inas­ tions on some of the botanical productions much as 'St. John and Fosberg did not find it of the Sandwich Islands. Mag. Nat . Hist. in 1934 after a careful search of the few re­ 5: 255-261. maining 'acres of native forest (personal com­ BENTHAM, GEORGE. 1870. Flora Australiensis. munication from Dr. St. John). Vol. 5. viii+599 pp. 1. Reeve & Co., This variety is named in honor of Dr. Carl London. Skottsberg, who for many years has been a student of Myoporum (and numerous other BRIGHAM, WILLIAM T. 1908. The ancient Ha­ genera) in the Pacific islands. waiian house. Bernice P. Bishop Mus., Mem. 2(3): iv+185-378, pls. 18-40, 178 DOUBTFUL AND EXCLUDED SPECIES figs. . Myopor~m ? euphrasioides Hook. & Am., Bot. BROWN, FOREST B. H. 1935. Flora of South­ Beechey Voy. 67, 1832. eastern Polynesia-III. Dicotyledons. Bernice This plant, described from Whitsunday P. Bishop Mus., Bul. 130: ii+386 pp., Island (= Pinaki 1., Tuamotus), is now known 9 pls., 70 figs. as Nesogenes euphrasioides (Hook. & Arn.) A. DC. Because Hooker and Arnott included the CHEESEMAN, T. F. 1903. The flora of Raro­ Tuamotus in the Society Islands, a number of tonga, the chief island of the Cook Group. writers listed Myopo rum euphrasioides from the Linn. Soc. Lond., Trans. Bot. II, 6(6) : 261­ latter group. 313, map, pls, 31-35. Polynesian Speciesof Myoporum-:WEBsTER 77

D E CANDOLLE, ALPHONSE. 1847. Prodromus mand of Captain F. W. Beechey .. . in the systematis naturalis regni vegetabilis. Vol. 11: years 1825, 26, 27, and 28. ii+ 485 pp. , 99 1-736. Victoris Masson, Paris. pls, Treuttel, Wurz, Treu ttelJr., & Richter, London. D EGENER, OTTO. 1930. Illustrated guide to the more common or noteworthy ferns and flowering KRAENZLIN, F. 1929. Beitrage zur Kenntnis plants of Hawaii National Park with descrip­ der familie der M yoporinae R. Br. Fedde tions of ancient Hawaiian customs and an Repert.Sp. Nov.Beih. 54: 1-129. introduction to thegeologichistory oftheislands. LEVEILLE, H . 1912. D ecades plantarum xv+ 312 pp ., frontisp. , 95 pls., 45 figs. novarum XC-XCII. Fedde Repert. Sp. Nov. Honolulu Star-Bulletin, Ltd. , Honolulu. 11: 63-67. EGLER, FRANK E. 1947. Arid southeast Oahu NADEAUD,). 1897. Note sur quelques plantes vegetation, Hawaii . Ecol. Monog. 17: 383­ .rares ou peu connues de Tahiti. Jour. Bot. 435, 41 figs. Morot 11(7) : 113-120. FORBES, CHARLESN. 1913. Notes on th e flora ROCK, JOSEPH F. 1913. The indigenous trees of of Kahoolawe and Molokini. Bernice P. . the Hawaiian Islands. iv+518 pp ., 215 pls , Bishop Mus., Occas. Papers 5(3) : 85-97, Honolulu. 7 figs. ST. JOHN, HAROLD. 1940. Itinerary of Hugh FORSTER, G . ( =). G . A.). 1786. Florulae in­ Cuming in Polynesia. Bernice P. Bishop suIarum australium prodromus. 103 pp . Mus., Occas. Papers 26(4): 81-90, portr. Dietrich, Gottingen, SKOTTSBERG, CARL. 1933. Myoporum in Raro­ FOSBERG, F. RAYMOND. 1943. The Polynesian tonga. Acta Horti Gotob. 8: 147-167, 48 figs. species of Hedyotis (Rubiaceae). Bernice P. Bishop Mus., Bul. 174: 1-102, 4 pls., 9 figs. SOLEREDER, HANS. 1899. Systematische Anato­ mieder Dicotyledonen. xii+ 984 pp. , 189 figs . GRAY, ASA. 1866. Characters of some new or Ferdinand Enke, Stuttgart. obscure species of plants, ofmonopetalous WETTSTEIN, R.VON. 1895. M yoporaceae. orders, in the collection of the United In: Engler & Prantl, Naturl. Pflanzenfam. States South Pacific Exploring Expedition . IV: 3b., 354-360, figs. 142-144. Wilhelm under Captain Charles Wilkes, U.S.N. Engelmann, Leipzig. Amer. A cad. A rts and Sci. , Proc. 6: 37- 55. WILDER, GERRIT PARMILE. 1931. Flora of HOOKER, WILLIAM JACKSON, and G .A. ofRarotonga. Bernice P. Bishop Mus., Bul. ARNOTT. 1830- 1841. The botany ofCaptain 86: 1-113, 8 pls., 3 figs. Beecbey's voyage comprising an account of the plants collected by Messrs. Lay and Collie, and ZIMMERMAN, ELWOOD C. 1948. Insects ofHa­ other office rs of the expedition during the voyage waii. Vol. I : Introduction. xx+ 206 pp., to the Pacific and Bering's Straits,performed in 52 figs. University of Hawaii Press, Ho­ His Majesty's ship, Blossom, under the com- nolulu.