A New from Kauai, Hawaiian Islands

BENJAMIN C. STONE and IRWIN LANE1

THE MOST RECENT revision of the Hawaiian Polynesiotis, with 19 , principally de­ species of Hedyotis () is that of F. R. veloped in Hawaii, with a secondary center Fosberg (1943), who treated all the Polyne­ in southern Polynesia . This subgenus has five sian species, describing many new forms . The sections: Wiegmannia, Protolsadua, Gouldiopsis, as considered by Fosb erg includes, with Bikkiocarpa, and Austrogouldia. good reason , the segregate genera The new species described herein fits well 1., Gerontogea Cham . and Schlecht., Kadtta into the subgenus Polynesiotis and into the Cham. and Schlecht., Diplophragma Meisn., Wiegmannia. However, it differs in as well as Gouldi« romanzoffiensis A. Gray, a two of the key characters employed in Fos­ species not properly in Gouldi«. berg's key to the subgenera. First, in our The type species ofHedyotis is H. auricalaria specimens the stigmas are consistently quadri­ 1. , which forms also the basis ofthe subgenus fid, not bifid; second, the width of the corolla Hedyotis. This subgenus contains the species tube is not "much less than 1'3 the length" of native to southern Asia which have axillary the corolla tube, as the key states , bu t is in inflorescences, indehiscent or septicidal fruits, some cases as wide as it is long. At first and usually a depressed habit of spermacoc­ glance the inflorescences seem axillary, but coid appearance. The sections and subgenera on closer inspection it can be seen that the y found in Polynesia are as follows. are strictly terminal; however, the first axillary bud below the inflorescence grows into a Subgenera: Oldenlandia, including only stem (or in some cases both of the two axil­ Hedyotisbiflora1., a wide-spread oftropi­ lary bud s), and these in turn eventu ally ter­ cal Asia, the islands of the Indian Ocean, minate in inflorescences; this gives the Malaysia, Micronesia, Melanesia, Fiji, and, an aspect ofbranching which might be termed in Polynesia, Samoa and Tonga. subscorpioid. Diplophragma, of southern and western Pol­ Within the section Wiegmannia of the sub ­ ynesia, eastern Melanesia, and Mi cronesia, genus Polynesiotis, our specimens are evidently but not known from Hawaii. very closely related to Hedyotis llttoralis (H bd.) , two variable species confined to the Fosberg, a striking and characteristic species ------Hawaiian Islands: - - distinguished by its fleshy leaves, which are closely set on a decumbent corky stem, and Oceanica, comprising the single species by its habitat on cliffs or rocks close to the transferred from Gouldia, and present onl y in sea. It is know n mostly from Molokai and southern and central Polynesia. M aui; although there exist specimens from

1 D epartment, University of H awaii. Man­ K auai, Oahu , and Hawaii, they are old (Hille­ uscript received March 4, 1957. brand's from Kauai and Oahu, Abbe Faurie's

139 140 PACIFIC SCIENCE, Vol. XII, April, 1958

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from Hawaii), and there have been apparently In the key to the species of section Wieg­ no other collections in these localities since mannia, insert in place of the lead to H . lit­ the original specimens were taken. Fosberg toralis: "2. Leaves, bracts , calyx lobes, and says "the plant has become quite rare, except ovary fleshy; leaves mainly sub -basal 3. on the windward coast of east Molokai. I 3. Corollas white, th e tube two or three searched for it without avail at Hanalei, times longer than wide; fruits subglobose, Kauai, in 1935. It has no t been found on crowned by th e persistent enlarged calyx Oahu since H illebrand's time." ...... H. littoralis Our specimens exhibit many features char­ acteristic of H. littoralis, including its habitat 3. Corollas green , the tube as long as wide preference, bu t in flower color, corolla shape, or up to twice as long as wide ; fruits and fruit shape they show clear-cut differences. usually conspicuously flattened, the per­ The general aspect of both species is very sistent enlarged calyx lobes spreadi ng similar, and the two are both quite different from the equatorial plane of the frui ts . . . in this regard from the majority of the other ...... H. St.-]ohnii" Hawaiian species of Hedyotis. The following changes in the keys should DES CRIPTION be made toaccomodate this new species. In He dyotis St.-Johnii B. C. Stone and 1. Lane '. the key ro the subgenera (Fosberg, 1943: 19), sp. nov. (Subgen. POLYNESIOTIS sect. under the second number 2, read: " Corolla WIEGMANNIA) fleshy or at least thickened, salverforrn, an­ thers included or the tips barely exserted , Suffrutex parvus non- vel pauci-ramosus style shorter than or subequ al with tube, bifid decumbens in scopuloris saxatilis maritimis or quadrifid (lobes may cohere), seeds an­ Kauaiensibus habitans, caulis lignosus longi­ gular. . ~.. . . 3." Under the second 3, read: tu dinaliter sulcatus, petiolis connatis et fasciis " Inflorescence terminal, terminal and axillary, suberos is inter foliosis ambis cinctis, folia or seemingly axillary bu t terminating the opposita dense conferta irregulariter ellip ­ main stem, the branch below arising from the tico-acuminata ad apicem saepe asymetrice first axillary bud (s) ...... 4." Under the sec­ curvata in vivo subcarnosa convexa supra ond 4, read: "Width of corolla tube usuall y fusco-viridia et nitida infra subglauca pullo­ much less than Y3 the length, but in some as venosa, bases foliorum lati-petiolatae vel ala­ wide as long; cyme usually many-flowered; tae bases bini connatae, laminae 5-1 4 cm. fruit dry, or if fleshy, calyx lobes much over longae 2-5 .5 cm. latae marginibus integribus, 1 mm. long Polynesiotis." superficies foliorum (et bracteorum er cali­ In the key to the sections of Polynesiotis corum) minute albo -scabratus, costa nervique (Fosberg 1943: 23), under the first 2, read: levirer prominulentes, nervi laterales 4- 10 " Calyx lobes foliaceous, longer than hypan­ arcuatim adscendentes vix anastomosantes thium in flower, conspicuously accrescent in (rete venularum conspicuum sed min ut um), fruit, distinctly nerved, subscabrous in some; folia emortui persistentes vestirentes, inflor­ corolla limb not quadrangular in bud, in- escentia stricte terminalis pauci -ramosa usque florescence usually quite glaucous . 17 cm. longa remote bracteata thyrsoidea tri­ Wiegmannia." partite cymosa , ramus quisque dichasius fere

FIG. 1. A-F, Hedyotis littoralis. A-I, Faurie 374, Mol okai; 1, calyces, l a, fruit ; 2,Faurie 375, H alawa, Hawaii, fruits . B, Skottsberg 800, East Maui, 250 m. alt. ; 1, calyx, young ; l a, fruit. C, Degener and Nitta 9430, Wailau, Molokai; 1, bud; l a, fruit. D , same, 2d sheet ; 1, buds and young calyces; la, fruit . E, Forbes 237-M, Keaneae , East Maui; 1, flower; la, fruits. F, leaves, 1, 2, Rock 7003, Wailau, Molokai; 3, Faurie 374; 4, Fosberg 13451, Wailau, Molokai; 5, Forbes 237-M ; 6, Skorrsberg 800; 7,D egener and Nitta 9430, sheet 2. (All in Bishop M us.) All natural size. 142 PACIFIC SCIENCE, Vol. XII, April, 1958

l cm

FIG. 2. Hedyotis St.-Johnii. a, Young flower; b, bud showing calyx; c, cross -section of bud, show ing young coroll a; d, top view of bud with calyx lobes spread; e, anther, front and side; fl , young pistil; f2, older pistil; g, fruit, top view, showing calyx expans ion and deh iscence lines; h, seeds. All X 7Y2. 143 New Hedyotis-STONE AND LANE •

infl.

3

2

1 o em

c

infl.

FI G. 3. A, Hedyotis St.-Johnii, habit sketch,(from Kodachrome by B. C. Scone); B, stem, branching pattern, showing rerminal inflorescence s; C, inflorescence; D, representarive leaves. A- X Y3; B-D, X 1. 144 PACIFIC SCIENCE, Vol. XII, April, 1958

11 floris ferrens, flos omnis bracteolo unico em. wide, the surfaces minutely white-scab­ subulato fere 1 mm. longo subtenrus, brae­ rous, the leaves when living rather fleshy, teae 2 rare 4-6 ovatae vel subcordatae ad convex, dark shining green above and rather apicem acutae vel subacuminatae 1-3 em. glaucous, but with darker veins, below; veins longae 1.3-2.1 em. latae inferiores longiores, barely prominent, the 4-10 lateral nerves curv­ flos junior subcuboideus, flos vetustior urceo­ ing parallel, hardly anastomosing; the vein­ latus 8-11 mm. diam., pedicellus 2 mm. lon­ reticulae apparent but delicate. Inflorescence gus, calyx in alabastro valvatus 4-lobatus, lobi terminal but seemingly axillary because of calycis hastati subrhomboidali 4-6 mm. longi subscorpioid branching, the inflorescence up quam corollam longiori, corolla in alabastro to 17 em. long, a thyrsoid compound tripar­ ad apicem depressa 5-8 mm. longa lobi in tite cyme, with each branch a strict dichasium flore leviter divergentes 1.5-2.5 mm . longi ofabout 11 flowers, each flower subtended by viridi, stamina subsessilia cum corolla adnata a bracteole 1 mm. long, subulate in form; the vix exserta, antherae 1.5 mm. X 0.8 mm. , pistil dichasial branches average 1.5 em. in length; 1, ovarium inferius, hypanthio discoideo, the entire cluster of flowers may reach 2-3 stylus ca. 3 mm. longus in basi pubescenti ad em. in width. Flowers in bud subcuboid, at apicem quadrifidem, fructus siccus lentifor­ anthesis urceolate, when mature 8-11 mm. mis vel subglobosus calyx persistens accre­ in diameter; calyx glaucous green, glabrous scens ad mediam fructus cinctus, fructus 4­ except for minute white scabrae, in bud val­ locularis dehiscens primo loculicidalis illo vate, subcuboid, with four hasrate subrhom­ tempore septicidalis, maturitas 4-6 mm. boidallobes whose basal corners are recurved­ diam., semina angulosa nigra 0.2 mm. longa. approximate, free; calyx longer than the Ab. H. littoralio corolla brevioribus urceo­ corolla ; the calyx-lobes in bud ca. 2 mm. lati viridi (non albi) et fructu lentiformi long, in flower ca. 3 mm . long, in fruit ca. differt. 8-10 mm .long; corolla green, in bud valvate, Suffrutescent, decumbent, sparsely branch­ the four lobes inflexed at the top of the bud ing plants growing on rocky cliffs near the and forming a depression; in flower spreading sea (known only from near Kalalau, Kauai), or barely ascending, darker green within; with woody stems up to 30 em. long covered corolla in bud 2-3 mm. long, in flower 5-8 by a corky, longitudinally grooved epidermis mm. long, the lobes 1.5-2.5 mm. long, sub­ and banded by corky rings and the connate triangular, the neck constricted, the outer bases of the petioles, the stems usually hidden edges of the petals somewhat recurved or by the congested persistent opposite dead and appearingridged. Stamens 4, subsessile, adnate living leaves, the branching pattern subscor­ to the constricted neck of the corolla, the tips pioid and the branches somewhat parallel, barely exserted, anthers 1.5 X 0.8 mm., the inflorescences terminating each branch and a connective-filament 0.1-0.2 mm. long, an­ new branch growing from the axillary bud ther-cells 4. Pistil 1, the ovary inferior, the just below the inflorescence (or sometimes hypanthium discoid, in flower not squared both buds developing into branches), these but becoming slightly so in fruit, 3-4 mm. branches in turn terminating in inflorescences, wide in flower; style deeply quadrifid, the 4 etc. The stems may reach a diameter of 1-2 lobes cohering; basal part of the style pu­ em. at the base, and are light brown to whitish bescent; lobes of the style 0.4 X 0.15 mm. in color. Leaves simple, entire, elliptical­ Fruits dry, flattened-lentiform to depressed­ acuminate with assymmetric curved apices, subglobose, the calyx-lobes persisting-accres ­ the length-width ratio rather variable, the cent to about twice the size they are in flower, petioles broad or alate and conspicuously lobes fenestrate, 5-7+nerved, spreading, connate, the blades 5-1 4 em. long and 2-5.5 attached at the equator of the fruit; fruit New Hedyotis-STONE AND LANE 145

dehiscing first loculicidally across the disc, only two collections of H. St.-Johnii are from the pyrenes later separating by a septicidal nearly the same locality along the Napali slit. Seeds angular, blackish, papillose to gra­ coast (Honopu and Kalalau are only about nulate, small, ca. 0.2 mm . long. 3 miles apart, as the crow flies), and Hille­ HOLOTYPE : Hawaiian Islands, Kauai: Be­ brand 's specimen was from Hanalei, perhaps tween Kalalau and Honopu, cliffs at end of 15 miles away and in a less rugged and rocky beach; plants growing on rocky ledges and area, the possibility still exists that this latter in crevices dashed by ocean spray, 10-30 ft. specimen represents H . littoralis. elevation, in association with Artemisia am­ Hedyotis St.-Johniiwas first collected in 1947, tralis and Lipochaeta succulenta, December 24, by St. John, Britten, and Cowan , and was 1956, Benjamin C. Stone no. 1470 (2 sheets, rediscovered in 1956, during a trip into the in Bishop Museum).(This locality is actually isolated and remote Kalalau Valley led by the same as the following.) Harold St. John. About 10 plants were seen, SPECIMENS EXAMINED : Hawaiian Islands, at various heights from 10 to 30 feet up, on Kauai: Third gulch from east end of Honopu the vertical basalt faces of the cliffs at the (Kalalau Trail), rare, in crevices of vertical west end of the beach between Kalalau Valley basalt sea cliff, 10 ft. alt., herbaceous, leaves and Honopu Valley. Flowers and fruit seemed fleshy, above dark shiny green, below pale abundant, and the colony though small green with darker veins ; infl. green, the buds seemed healthy. The plants were continuously cuboid, the calyx lobes valvate, within pale sprayed by a fine spume from the breaking green, the corolla lobes valvate, within dark waves. The only plant associates were Arte- green, without pale green, assurgent; anthers .misia australis Less. and Lipochaeta succulenta exserted, yellow ; December 31, 1947, Harold DC. , though other species were growing St. John, E. J. Britten, and R. S. Cowan no. nearby at the base of the cliffs back of the 23,207 (2 sheets , in Bishop Museum). beach. DISCUSSION ACKNOWLEDGEMENTS Because ofthe close vegetative similarity of The authors would like to express their the species Hedyotis littoralis (Hillebrand) Fos­ appreciation to Harold St. John, Senior Pro­ berg and the newly described H. St.-Johnii fessor of Botany at the University of Hawaii, Stone and Lane, there is the possibility that in whose honor this new species is named, Hillebrand's collection from Hanalei, Kauai, for his generous assistance in: the preparation might represent H. St.-Johnii rather than H. of this paper, and for the opportunity and littoralis. Hillebrand (1888) added after the inspiration to study the Hawaiian flora; and description of his H. littoralis, " On rocks near to Mrs. Nancy Higgins, without whose in­ the seashore in Waikolu, Molokai! and Hana­ genuity the second collection might never lei, Kauai! A single damaged corolla only was have been made. available for examination; the position of the anthers, shape of corolla lobes, and relative REFERENCES length of style remain therefore doubtful." FOSBERG, F. RAYMOND. 1943. The Polynesian Possibly also the color of the corolla in the species of Hedyotis (Rubiaceae). Bernice P. Hanalei specimen was unknown. At any rate, Bishop Mus., Bul. 174: 1-101. until new and definite collections of H. lit­ toralis are made on Kauai, it must remain a HILLEBRAND, WILLIAM. 1888. Flora ofthe Ha­ matter of doubt whether both of these species waiian Islands. Heidelberg. I-XCIV + 1­ are actually present there. Since, however, the 673 pp. Cf. p. 167.