Monograph of the Hawaiian Species of ()l Hawaiian Studies 34 HAROLD ST. JOHN 2

THE GENUS Gouania was early noted as a part Gouania in the Hawaiian Islands, up to the of the Hawaiian flora by F. J. F. Meyen (1835), present. and he described a new species, basing it upon There is, however, an issue as to the generic his own collection made on Diamond Head, placement of the local species that needs discus­ Oahu, in 1831. He published the species in his sion. In the first edition of Engler and Prantl's narrative (1835:150) as G. integrifolia, but Pflanzenfamilien (1896:424-425) Weberbauer this epithet was a homonym of the earlier one published the new genus Pleuranthodes to in­ for another species by Lamarck. When Walpers clude only the two Hawaiian species, P. orbicu­ prepared the botanical report on the collections lare (Walp.) Weberb. and P. Hillebrandti of this voyage of the "Prinzess Louise," he (Oliver) Weberb. As printed, both of these realized that Meyen's binomial was unavailable, epithets were incorrect. As indicated above, the so published it anew (1843:323) as Gossania earlier and valid epithet for the first species is orbicularis. There was no existing genus Gos­ Meyeni, and that for the second is Hillebrandi. sania, and it is evident that no new genus was This latter confusion was due to there being intended. The generic name was apparently the two German botanists, a more prominent and printer's attempt to reproduce the author's active one, Johann Maria Hildebrandt, and a handwriting, and it was not corrected in proof­ lesser known amateur, William Hillebrand, a reading. It is apparent that Walpers intended to physician long resident in Hawaii, and author publish the binomial Gouania orbicularis. It was of the standard flora of the Hawaiian Islands. so listed in the Index Kewensis, and this render­ The second species was named in honor of the ing seems to be correct. However, this second latter man, and his name contains no letter t. binomial was unnecessary, as in 1840 Steude1 Weberbauer treated Gouania as also present had already published for it the valid name in Hawaii, and consisting of 30-40 species of Gouania Meyeni (Steudel, 1840:703). the tropics of the world. He diagnosed the genus A second species was collected in 1840 by as having the style 3-lobed; disk mostly produced C. Pickering and W. D. Brackenridge, the into one lobe in front of each calyx lobe; fruit botanists of the Wilkes Expedition, in the 3-celled, 3-winged; inflorescence a spike or Waianae district, and published as G. vitifolia panicle; leaves entire or toothed; , or, if Gray (1854:283). , the fruit wingless. Pleuranthodes was There were some other early collections of the said to have the style 2-lobed; disk with 5 short genus, but no additional species were published broad lobes in front of the calyx lobes; fruit until 1888 when W. Hillebrand's Flora of the 2-4-celled, 2-4-winged; inflorescence a cyme; leaves entire; erect shrubs. Hawaiian Islands was printed. In it he added (1888:83) two new species, G. Bishopii Hbd., In the second edition of this work (Engler from Lahaina, Maui, and G. Hillebrandi Oliver, and Prantl, 1953:166-171) Suessenguth es­ from Kula and Lahaina, Maui. A fifth species, sentially kept the same classification. He retained G. Fauriei (LevI.) St. John, was discovered on the invalid epithet Pleuranthodes orbiculare Molokai in 1910 and published as Isodendron (Walp.) Weberbauer, but corrected that of the Fattriei by Leveille (1912 :63). That summa­ second species to P. hillebrandii (Oliver) We­ rizes the full story of the accepted species of berbauer. He concluded with the statement, "Beide Arten vielleicht durch Zwischenformen verbunden." That is: "Both species perhaps are 1 Manuscript received January 23, 1969. 2 B. P. Bishop Museum, Honolulu, Hawaii 96818, connected by intermediate forms." There were U. S. A. no specimens of either species in his Muenchen 507 508 PACIFIC SCIENCE, Vol. XXIII, October 1969 herbarium, and he cited no evidence to support genus G01/ania evidences what has happened to his depreciation of the two species. Nor did he the native flora of the dry lowland regions; it study and annotate any specimens of these has suffered an almost complete extinction. How species from other herbaria. His statement was many other native of the lowland region wholly gratuitous. Of the genus GOf/ania, he were exterminated before they were collected estimated there were 66 species. and recognized, we cannot even guess. In review of the stated diagnostic characters Significant morphological characters in the between Gottania and Plettranthodes, there are local species of Gottania are found in the shape no significant differences in the lobes of the and nature of the fruit, in the seeds (known in style, in the disk (five lobes beyond the five only three of the species), in the inflorescence, calyx lobes is the same as one lobe beyond each in the sexuality of the flowers, in leaf form, calyx lobe), in the cells and wings of the fruit, size, and pubescence, and in the plant habit. or in the inflorescence (since we now can state Several of the species have all their flowers that Got/ania has panicles, spikes, or cymes, quite similar in appearance, but actually they while Plettranthodes has spikes or cymes) . There are polygamous or functionally dioecious. This are left only the items that Plef/ranthodes con­ is certainly so in G. Hillebrandi, G. Meyeni, tains erect shrubs with entire leaves, while G. Remyi, G. sandu;ichiana, and G. vitifolia. Of Gottania has climbing shrubs with tendrils or most of the other species, the inflorescences are erect shrubs, and the leaves are toothed or en­ too sparse or too immature to allow observation tire. Since the fundamental characters of flower of this state of sexuality. It is, however, quite and fruit are identical, and since the only likely that they also are polygamous. The polyg­ actual differences are overlapping ones of habit, amous condition is not universal, but is common the genus Plet/ranthodes Weberbauer is here in the species of GOftania in other tropical areas. reduced to the synonymy of Gottania Jacq. Carlquist has advanced the theory (1965 :267­ The Hawaiian species of GOftania are native 270) that genera that have migrated to and to the dryland scrub or the lower dry forest established themselves in island floras tend in Zone. This is the region most drastically affected their local speciation to develop polygamous or by human settlement and by the eating and tram­ dioecious flowers. In the Hawaiian flora he lists pling by introduced grazing animals. The vege­ the examples Pelea, M yrsine, Brottssaisia, and tation of these lowlands was largely destroyed others. The genus Gottania can now be added before it was observed or collected by scientists. to this list. The result of the same wholesale destruction was Seed production is rare in the Hawaiian observed by the writer in his revision of the species of Gottania. See our discussion of this in Hawaiian species of Isodmdrion (1952, p. the treatment of the extant species G. Hille­ 216). Most of the recent Hawaiian botanists brandi. have never found a living specimen of Gottania, When one surveys the local distribution of and of the 14 species only 3 have been found the species of Gottania, it is seen that the genus and collected since 1886. was present on all the large islands of the The four species known to Hillebrand (1888: Hawaiian group, except Kauai and Niihau. 82-84) are still considered good and are here Maui had 5 species, the largest number, Oahu 3, accepted. The present writer has found and Kahoolawe 2, Hawaii 2, Molokai 1, and Lanai collected G. Hillebrandi and brought it into 1. They were restricted to semiarid habitats, cultivation, and G. vitifolia was rediscovered by and on all the islands (except Lanai) were Russ in 1929 and by Degener in 1932, and it may well be extant. The writer's study of the found on the leeward lower slopes of the genus has been mostly upon early collections mountains (except for G. thinophila, which now found in the older herbaria. Based upon occurred on Maui on the sand dunes of the these collections he now publishes 10 new windward shore of the isthmus). Two of the species, bringing the total in Hawaii to 14 species, G. pilata and G. sandu;ichiana cannot species. Of these, 6 have not been collected be mapped precisely, as their locality data are since 1855, 3 not since 1865, 2 not since 1886, merely the names of the islands, Maui and and 1 not since 1910. Like Isodmdl'ion, this Hawaii. Hawaiian Species of GOllcmia-ST. JOHN 509

KEY TO THE SPECIES

A. with tendrils; blades toothed, B. Blades elliptic to ovate, subcordate 7. G. Bishopii B. Blades broadly elliptic, deeply cordate, C. Blades beneath appressed pilosulous on the principal veins 2. G. vitifolia C. Blades beneath densely shaggily pilosulous. 13. G. hawaiiensis A. Erect shrubs without tendrils; blades entire, D. Blades suborbicular, emarginate 1. G. Meyeni D. Blades narrower, not emarginate, E. Blade intervals below glabrous; blades shortly subacuminate obtuse; leafy branchlets glabrate in lower part. 3. G. Oliveri E. Blade intervals hairy below; leafy branchlets with persistent pilosity, F. Blades tapering to an acute apex, G. Blades lanceolate to narrowly so; petioles 15-30 mm long; calyx lobes 1.3 mm long; disk with one broad, evident, retuse lobe opposite each calyx lobe. 10. G. pilata G. Blades elliptic to elliptic-ovate; petioles 4-17 mm long; calyx lobes 1-1.1 mm long, H. Hypanthium semiorbicular; disk with one slight emarginate lobe opposite each calyx lobe; flowering spikes 5-10 mm long. 12. G. ettctlllata H. Hypanthium cupulate; disk with 2 prominent rounded lobes opposite each calyx lobe; flowering cymes 25-35 mm long. 11. G. Remyi F. Blades elliptic, not tapering to the apex, 1. Blades at maturity sparsely hairy beneath, J. Wings of capsule equally wide from base to tip; stipules caducous, K. Peduncle 6-17 mm long; petals 1.3 mm long; calyx lobes 1.5-2 mm long 6. G. Hillebrandi K. Peduncle 20-45 mm long; petals 1 mm long; calyx lobes 1.3 mm long. . 14. G. sandwichiana J. Wings of capsule broader at apex; stipules long persis- tent. 4. G. Fauriei 1. Blades densely appressed pilose beneath, L. Peduncle 10-40 mm long; branchlets half spread­ ing pilose, M. Blades 3-6.5 cm long, the lower surface hid­ den by the dense pilosity; capsules equally winged from base to tip..... 8. G. Lydgatei M. Blades 2.2-3.5 cm long, pilose below but the surface visible; capsule wings narrowed to the base. 9. G. thinophila L. Peduncle 2-4 mm long; branchlets subappressed stiff villous; blades 1.4-3.7 cm long. 5. G. Mannii 510 PACIFIC SCIENCE, Vol. XXIII, October 1969

1. Gouania lI1eyeni Steud., Nomencl. Pl., ed. the stamen; filaments 1.3 mm long, subulate; 2, 1:703, 1840. (Fig. 1) anthers 0.4 mm long, suborbicular; disk notched G. integrifolia Meyen, Reise 2: 150, 1835, to each filament base and shallow retuse between non Lam. (1789). each pair; styles 2 and 0.7 mm long; stigmas Gossania orbicularis Walp. in Meyen, Obs. apical, discoid; lateral flowers apparently func­ Bot., Nov. Acta Acad. Caes. Leop.-Carol. tionally staminate, with the calyx 2.5 mm long, Nat. Curios. 323, 1843, evidently a mis­ closely appressed ascending white puberulent print and intended to be Gouania orbicu­ without, the tube 0.8 mm long, hemispheric, laris Walp., and it was so treated in Ind. the lobes 2 mm long, lanceolate, ending in an Kew. acuminate tip 0.3-0.4 mm long, within glabrous and thickened on the margins and midrib; petals ORIGINAL DIAGNOSIS: "G. fruticosa, erecta, 0.7 mm long, white, clawed at base, obovate, simplex, foliis longe petiolatis coriaceis sub­ saccate and deeply cucullate and enclosing a rotundis retusis integerrimis subtus petiolisque stamen; filaments 0.3 mm long, subulate; anthers puberulis, racemulis axillaribus, paucifloris 0.2 mm long, didymous suborbieular; disk 1.5 petiolo brevioribus, calyce hirsuto. Sandwich Is., mm in diameter, notched at the base of each Oahu, Demant-Hilgels, June 29, 1831, F. J. F. filament; the style 0.3 mm long, bifid; ovary Meyen." evidently sterile. EXPANDED DESCRIPTION (from Wilkes Expl. HOLOTYPE: "Sandwich Is., Oahu, Demant­ Exped. coIL): , with curved ascending Hilgels, June 29, 1831, F. J. F. Meyen." The branches, at least 25 em tall; stem at base 3 holotypic specimen, once in the Berlin her­ mm in diameter, terete, brown, with longitudinal barium, was destroyed in World War II, and fissures and with remnants of pubescence; leafy no isotype has been found in any of the prin­ branchlets 1 mm in diameter, rather densely cipal herbaria of the world. Holotype not appressed pale puberulent; internodes 2-5 mm examined. long, straight; nodes with a bracket-like enlarge­ ment at the leaf scar; stipules 1-1.5 mm long, SPECIMENS EXAMINED: Sandwich Islands, lanceolate, appressed puberulent on the outer U.S. Exploring Expedition under Captain side, persisting nearly as long as the leaves, Wilkes (GH, us) . The expedition made a leaving an elliptic scar; petioles 5-19 mm long, lengthy stay in Honolulu harbor, and it is likely appressed ascending white puberulent; blades that the botanists visited the nearby, prominent, 1.3-2.9 em long, 1-2.5 em wide, mostly very Diamond Head. broadly elliptic, but a few of the apical and The epithet honors the collector of this largest ones suborbicular and subemarginate, species, Franz Julius Ferdinand Meyen (1804­ firm chartaceous, entire, the margin smooth, or 1840), surgeon and botanist on the world toward the apex full and slightly undulate, the voyage of the Prussian ship "Prinzess Louise." upper surface olive green and completely His real specialty was plant physiology, but he glabrous, the lower surface rather sparsely and did creditable exploring on this voyage. almost invisibly appressed white pilosulous; in­ florescences numerous, axillary, 5-10 mm long, 2. Gottania vitifolia Gray, U.S. Expl. Exped., cymose, densely appressed brownish pilosulous; under Wilkes, 15 (1 ): Botany, Pha­ peduncle 2-3.5 mm long; flowers 3-5; buds nerogamia 1:283, 1854. (Fig. 2) 1.5 mm long, greenish, obturbinate; the Degener, 0., and A. Greenwell, Fl. Haw., terminal perfect flowers with the calyx in fam. 215: text and fig. 7/1/47. anthesis 2.5-3.5 mm long, closely appressed EXPANDED DIAGNOSIS OF HOLOTYPE AND ascending white puberulent, the tube 1.5 mm ISOTYPE: Woody ; stems 4 or more mm in long, hemispheric, the 5 lobes 1.5-2 mm long, diameter, gray, with slight longitudinal ridges, lance-ovate, thick and coriaceous, within gla­ glabrate; branchlets 1-3 mm in diameter, closely brous and the slender tip with a raised triangular subappressed rusty puberulent; leaf scars reni­ thickening; petals 1.2 mm long, white, spatulate, form; nodes slightly swollen; internodes 15-80 concave and deeply cucullate hooded, enclosing mm long on the main stems, 3-10 mm long on Hawaiian Species of GOlfallia-ST. JOHN 511

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FIG. 1. Goual1ia Meyelli Steud., from Sandwich Is., U.S. Exploring Expedition. a, Habit, X 1; h, bud of terminal perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; f, bud of staminate flower, X 15; g, staminate flower, X 20. 512 PACIFIC SCIENCE, Vol. XXIII, October 1969

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FIG. 2. Gray, from holotype, Oahu, U.S. Exploring Expedition. a, Habit, X 1; b, Rower­ ing shoot, X 1; c, bud of perfect Rower, X 15; d, perfect Rower, X 15; e, petal, X 30; f, stamen, X 30; g, seed, lateral view, X 5; h, seed and hilum, apical view, X 5; i, pistillate Rower, X 15. Hawaiian Species of GOltania-ST. JOHN 513 axillary branchlets; tendrils unbranched, coiling, anae] U.S. Exploring Expedition, 1838-42 a single one produced at one of the lower nodes (us)! Isotypes (BISH, GH)! Types examined, of an inflorescence (hence mere leafy branches and the holotype is here illustrated. lack tendrils); stipules 2.5-4 mm long, firm, SPECIMENS EXAMINED: Hawaiian Islands, ovate, acute, appressed rusty pilose, caducous; Oahu, Keaau, Aug. 1929, G. W. Russ (BISH); petioles 3-14 mm long, densely subappressed first gully makai on south side of Keaau Valley, rusty puberulent; blades 30-63 mm long, 20­ in dry rocky Aleurites forest climbing to highest 46 mm wide, broadly ovate, the base deeply trees like grape vine, on verge cordate with a narrow sinus, the apex acute or of extinction, shortly subacuminate, texture firm chartaceous, Feb. 7, 1932, O. Degener, K. Park & Y. Nitta coarsely crenate, the apex of the teeth tipped 8,731 (G, s, us); ditto, same plant, March 23, with a hydathode, the upper surface dark green 1932, Degener, Park & Westgate 8,732 (BISH, and appressed soft puberulous, the lower sur­ G, us); ditto, June 1932, Degener, Dr. & Mrs. face whitish green but rusty soft puberulous Bilger 8,733 (us). only on the principal veins; inflorescences This is one of the two or three species that numerous, 1-7 cm long, borne on the short have survived until recent times, and may still axillary branchlets, densely rusty hirsutulous; be extant. flowers spicate in glomerules, these crowded The epithet was obviously derived from the above; buds 1.5 mm long, subglobose, densely Latin, vitis, a vine or grapevine, and folium, reddish brown hirsutulous; perfect flowers (or leaf, and its toothed leaves do bear a resemblance perhaps only functionally staminate, and it is to those of the grape. noted that Gray called the upper flowers sterile) with calyx tube 0.7 mm long, funnel­ 3. Gouania Oliveri sp. nov. (Fig. 3) form; calyx lobes 5 and 1 mm long, broadly DIAGNOSIS HOLOTYPI: Licet frutex, ramulis ovate, acute, concave, valvate, thick and sub­ 1.5-3 mm diametro mox glabratis subpurpuratis coriaceous, within with a thickened strip up the laevibus sed in sicco longitudinaliter sulcatis, middle, without densely appressed ascending ramulis novellis sparse adpresse adscendente hirsutulous; petals 0.8-1 mm long, 0.3 mm albi-pilosis, internodis 3-15 mm longis rectis, wide, spatulate, deeply cucullate, the margin and nodis subincrassatis, stipulis 2-4 mm longis apex infolded and enclosing the stamens; fila­ linearibus dense adpresse brunnei-pilosis, petiolis ments 1 mm long, subulate; anthers 0.3 mm 15-44 mm longis teretibus gracilibus sparse long, quadrate subglobose; disk concave, the adpresse adscendente albi-pilosis, laminis 3.2-6 margin 10-lobed; pistillate flowers similar but cm longis 1.6-3.8 cm latis chartaceis subtilibus calyx tube 0.7 mm long, funnelform; the 5 late ellipticis in basi rotundatis in apice sub· calyx lobes 0.8 mm long, ovate, acute, valvate; acuminatis obtusis marginibus integris supra petals 0.7 mm long; stamens 0.17 mm long, obscure viridibus glabris infra in midnervo slender subulate, erect, with a tiny abortive sparse adpresse albi-pilosis et minime in nervis anther; stigmas 3, erect, 0.2 mm long, linear; secundariis in pagina remote adpresse albi­ capsules 9-10 mm long, suborbicular, swollen pilosulis pilis adscendentibus, inflorescentiis 3-5 over the seeds, 2-3-winged, the wings 2.5-3 cm longis axillaribus racemosi-cymosis adpresse mm wide, running the full length of the cap­ adscendente albi-pilosulis 3-5-floriferis, pedun­ sule and of equal width throughout; seeds culis 3-4 cm longis gracilibus, pedicellis 1-4 3.5-3.8 mm long, 3-3.5 mm wide, 1.5-1.8 mm mm longis, alabastris 2-2.5 mm longis ovoideis thick, broadly ovoid, in cross section oblate acutis valde adpresse adscendente albi- vellutei­ trigonous, but the back convex, dark brown, pilosulis, lobis calycis 2-2.2 mm longis del­ smooth, very shiny. toideis vel anguste deltoideis intra glabris et in midnervo marginibus apiceque triangulari in­ EXPANDED DESCRIPTION: Petioles 3-28 mm crassatis extra sparse adpresse adscendente long; blades 30-110 mm long, 20-69 mm wide. pilosulis, petalis 1.4 mm longis 0.7 mm latis HOLOTYPE: "Sandwich Islands: on dry hills, spatulatis cucullatis cum marginibus apiceque in the district of Waianai, Oahu." [= Wai- involutis et staminem inclusis, filamentis 1 mm 514 PACIFIC SCIENCE, Vol. XXIII, October 1969

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FIG. 3. Gouania Oliveri St. John, from holotype, Oahu, Lihue, Hillebrand. a, Habit, X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; I, capsule, X 4; g, venation of blade, X 4; h, seed, lateral view, X 5. Hawaiian Species of GOllal1ia-ST. JOHN 515 longis late subulatis, antheris 0.5 mm longis SPECIMENS EXAMINED: [Hawaiian Islands], cordati-ovoideis, margini disci vadose undulati Oahu, Makaha, ex Mus. Bot. Berolin., [W. lO-lobati, capsulis 11-12 mm longis late el­ Hillebrand], (BISH). Sandwich Is., U.S. Explor­ lipticis vel suborbicularibus compressis bilobatis ing Expedition, (p). bialatis in basi cordatis in apice late retusis axili 9 mm longo, seminibus 5 mm longis 3 mm latis DISCUSSION: From the few meager old speci­ ellipticis compressis brunneis laevibus lucidis. mens available, it is not possible to determine the range of sexuality of the flowers. DIAGNOSIS OF HOLOTYPE: Evidently a shrub; This collection by Hillebrand formed part of branchlets 1.5-3 mm in diameter, early glabrate, his concept of the species G. orbicttlaris Walp. purplish, smooth, when dried with longitudinal which is now called G. Meyeni Steud., but the ridges; young branchlets remotely appressed Waianae Mountains collections are now found ascending white pilose; internodes 3-15 mm to be specifically distinct from that species. It long, straight; nodes very slightly enlarged has not been collected in the last hundred years, below the leaf scar; stipules 2-4 mm long, and appears to be extinct. linear, obscured by the brownish appressed The specific epithet is chosen to honor Daniel pilosity, leaving a rounded scar; petioles 15-44 Oliver (1830-1916) of Kew who assisted mm long, terete, slender, rather sparsely ap­ Hillebrand in interpreting the Hawaiian species pressed ascending white pilose; blades 3.2-6 of the genus. em long, 1.6-3.8 em wide, thin chartaceous, broadly elliptic, the base rounded, the apex 4. Gouania Fauriei (levI.) comb. nov. shortly subacuminate, obtuse, the margins entire, (Fig. 4) Isodendron (= Isodendrion) the upper surface dark green, glabrous, the FalJriei (levI., Fedde Repert. 11 :63, lower surface sparsely appressed white pilose on 1912. the midrib and even less so on the secondaries, the surface remotely appressed white puberulous, DIAGNOSIS HOLOTYPI: Licet frutex, ramis 2-6 the hairs ascending; inflorescences 3-5 em long, mm diametro teretibus deinde glabratis et cortice axillary, appressed ascending white pilosulous, griseo vel brunneo et squamato, ramulis novellis racemose-cymose, bearing 3-5 flowers; peduncle densiter subadpresse pilosis pilis albis vel rubri­ 3-4 em long, slender; pedicels 1-4 mm long; brunneis et persistentibus, internodis 2-7 mm buds 2-2.5 mm long, ovoid, acute, closely ap­ longis rectis, nodis incrassatis et ad cicatricem pressed ascending white to yellowish pilosulous; projectentibus, stipulis 3-5 mm longis lancei­ calyx lobes 2-2.2 mm long, deltoid or narrowly linearibus densiter adpresse pilosis persistentibus, so, within glabrous, the midrib, margins, and petiolis 10-20 mm longis teretibus gracilibus apical triangle thickened, without sparsely ap­ dense pilosis pilis in parte adpresse adscendenti­ pressed ascending pilosulous; petals 1.4 mm bus in parte subdivergentibus, laminis 2.3-3.8 long, 0.7 mm wide, spatulate, the margins and em longis 1-2.2 em latis subtiliter coriaceis el­ apex inrolled and enclosing the stamen; fila­ lipticis in basi rotundatis in apice subacutis ments 1 mm long, broad subulate; anthers 0.5 plerumque mucronatis marginibus integris supra mm long, cordate-ovoid; disk undulate, shal­ obscure viridibus glabris infra sparse adpresse lowly 10-lobed; capsules (4 seen) 2-lobed and albi-pilosulis midnervo et nervis secundariis cum winged, 11-12 mm long, broadly elliptic to pilis pluribus, inflorescentiis 1.5-2.5 em longis suborbicular, compressed, at base narrowly axillaribus racemosis subadpresse albi-brevi-pil­ cordate, at apex broadly retuse, the center axis osulis 2-5-floriferis, pedunculo 9-10 mm longo, 9 mm long; seeds 5 mm long, 3 mm wide, pedicellis 5-6 mm longis, alabastris 1.3 mm elliptic, flat, brown, smooth, shining. longis ovoidei-suborbicularibus densiter adpresse adscendente brunnei-pilosulis, lobis calycis 1.5 HOLOTYPUS: Hawaiian Islands, Oahu, Wai­ mm longis subovati-deltoideis apiculatis intra anae Range, near Lihue, June '61 [W. Hille­ glabris et midnervo marginibus apiceque incras­ brand], (BISH). Isotypes (BISH), and Waianae satis apice umbonato extra hirusutulis pilis 0.1 Range, received July 1865, Dr. Hillebrand 126 nun longis, petalis 1 mm longis 0.5 mm latis (K)! spatulatis cucullatis marginibus apiceque involu- 516 PACIFIC SCIENCE, Vol. XXIII, October 1969

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FIG. 4. Gouallia Faul'iei (LevI.) St. John, from Molokai, Makolelau, Forbes 123.Mo. a, Habit, X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30. Hawaiian Species of Gottania-ST. JOHN 517

tis staminem inclusis, filamentis 0.7 mm longis Molokai, Makolelau, July 1912, C. N. Forbes subulatis, antheris 0.25 mm longis thecis ellip­ 123. Mo. (BISH). Isotypes (ue, us). tici-oblongis in basi divergentibus, disco cum Isodendl'ion Fattl'iei LevI. was one of the 10 lobis rotundatis, stylis 3 et 0.7 mm longis many new Hawaiian species ill-conceived by parte Yz infera connatis, capsulis 8 mm longis Leveille. His diagnosis is 2;4 lines long, and suborbicularibus cuneatis sparse adpresse breve contains five descriptive words, phrased as a albi-pilosulis 3-lateratis 3-alatis, alis 1-1.5 mm contrast with I. pyrifolium Gray, a member of latis subsymmetricis sed in basi auriculatis. the Violaceae. There are two excellent isotypes of I. Fauriei, Faurie 693, from Kamolo, Molo­ DIAGNOSIS OF HOLOTYPE: Evidently a shrub; kai, in the Bishop Museum. It did represent a branchlets 2-6 mm in diameter, terete, finally new species, but, as was often the case with glabrate and with gray or brown checkered bark; Leveille, he placed it in the wrong genus, and young branchlets densely subappressed pilose, this time in the wrong family. the hairs white to reddish brown, long persistent; As shown by the original labels, Leveille internodes 2-7 mm long, straight; nodes en· had difficulty in placing this plant, and he first larged and bracket-like beneath the leaf scars; put it in Chrysophyllttm, then changed his stipules 3-5 mm long, lance-linear, densely ap­ mind, and described it as a new Isodendrion. pressed pilose, persistent, usually well outlasting Rock (1914:355), when reviewing Leveille's the leaves, leaving a rounded scar; petioles 10­ publications on Hawaiian plants, reduced I. 20 mm long, terete, slender, densely pilose, the Fauriei, but Rock, too, made a misidentification hairs partly appressed ascending, partly half in referring it to I. pyrifolium (as Isiodendron). spreading; blades 2.3-3.8 em long, 1-2.2 em St. John (1952:255), when monographing wide, thin coriaceous, elliptic, the base rounded, Isodendrion, referred I. Fattriei to the synonymy the apex subacute, mostly mucronate, the mar­ of Gouania Hillebrandi Oliver. gins entire, upper surface dark green, glabrous, The epithet here accepted honors the mis­ lower surface sparsely appressed white pilosu­ sionary and botanist Urbain Faurie (1846­ lous, the hairs ascending, the midrib and 1915). secondaries more densely appressed pilose; in­ florescences 1.5-2.5 em long, axillary, racemose, 5. Gouania Mannii sp. nov. (Fig. 5) half appressed white short pilosulous, bearing DIAGNOSIS HOLOTYPI: Licet frutex, ramis cum 2-5 polygamous flowers; peduncle 9-10 mm cortice brunneo squamato, ramulis foliosis sub­ long; pedicels 5-6 mm long; buds 1.3 mm long, adpresse brunneis rigidis villosis, internodis ovoid·suborbicular, densely appressed ascending 3-19 mm longis rectis, nodis paene incrassatis, brownish pilosulous; apical perfect flower with stipulis 3-5 mm longis anguste lineari-Ianceola­ its calyx lobes 1.5 mm long, subovate-deltoid, tis et densiter adpresse villosis et cum foliis apiculate, within glabrous and the midrib and persistentibus, petiolis 7-17 mm longis gracili­ margins thickened, merging into the much bus et subadpresse adscendente rigidi-villosis, thickened tip, triangular and with a projecting laminis 1.4-3.7 em longis 1.1-2.2 em latis sub· umbo, without appressed hirsutulous, the hairs coriaceis ellipticis vel obovatis late obtusis mar· 0.1 mm long; petals 1 mm long, 0.5 mm wide, ginibus integris supra olivaceis glabris infra ad­ spatulate, the margins and apex inrolled, cucul­ presse albi-pilosis et in nervis principalibus ad· late, enclosing the stamen; capsule 8 mm long, presse brunnei-villosis, inflorescentiis 3-6 mm cuneate suborbicular, sparsely white appressed longis axillaribus spicatis (vel cymosis ?) , short pilosulous, 3-sided, and the 3 wings sub­ pedunculo 2-4 mm longo densiter adpresse equally developed, 1-1.5 mm wide from base rigide brunnei-villosis, floribus 2-5 congregatis, to tip and at base downward auriculate. alabastris ovoidei-subglobosis, calycibus in flore 3 mm longis extra densiter adpresse adscendente HOLOTYPE: Molokai, Kamolo [ = Kamalo], brunnei-hirsutulis lobis 1.2 mm longis deltoideis 1,000 malt., junio 1910, Fattrie 693 (BISH), ovatis crassis, petalis 0.4 mm longis 0.3 mm isotype of Isodendl'ion Fattriei LevI. latis spatulatis profunde cucullatis et staminem SPECIMENS EXAMINED: Hawaiian Islands, inclusis, filamentis 0.4 mm longis lanceolatis 518 PACIFIC SCIENCE, Vol. XXIII, October 1969

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FIG. 5. Gouania Mannii St. John, from holotype, Lanai, Mann & Brigham 344. tI, Ilabit, X 1; h, bud of perfect flower, X 15; C, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; t, immature capsule, X 4. Hawaiian Species of GOllania-ST. JOHN 519 attenuatis, antheris 0.25 mm longis obcuneati­ The new epithet is chosen to honor Horace oblongis retusis, margini disci undulati et ad Mann, Jr. (1844-1866), disciple of Professor lobis calycorum bilobati, stylis 2-3 et 0.6 mm Asa Gray, and early explorer of the Hawaiian longis, capsulis immaturis 4-5 mm longis ad­ flora, and the author who began the first pub­ presse pilosulis orbicularibus sed acute trigonis lished flora of the islands. et 3-alatis, alis symmetricis. 6. Gouania Hillebrandi Oliver, in Hillebrand, DIAGNOSIS OF HOLOTYPE: Probably a shrub; Fl. Haw. Is., 83, 1888. (Figs. 6, 9g-j, branches with brown flaky bark; leafy branches 16a, b) shiny brown, subappressed stiff villous; inter­ Plellranthodes Hillebrandti (Oliver) We­ nodes 3-19 mm long, straight; nodes scarcely berb., in Engler and Prantl, Nat. Pflan­ enlarged; stipules 3-5 mm long, narrowly linear­ zenfam. 3(5) :424, 1897 (Sphalm.). lanceolate, concealed by the dense appressed P. hillebrandii (Oliver) Weberb. emend. similar villosity, persistent almost as long as the Suessenguth, Engler and Prantl, Nat. leaves; petioles 7-17 mm long, slender, con­ Pflanzenfam. ed. 2, 20d:166, 1953. cealed by the brownish stiff subappressed as­ cending villosity; blades 1.4-3.7 em long, 1.1­ REVISED DIAGNOSIS OF LECTOTYPE: "Shrub; 2.2 em wide, subcoriaceous, elliptic to obovate, the slender branches with a rust- or ash-colored the apex broadly obtuse, appearing mucronate, pubescence. Leaves ovate-oblong 1yz-2yz X but this appearance is from the tuft of villosity 14-1 inch, on petioles of yz-1 inch, obtusely from the apex of the midrib, the margins entire, pointed, entire, dark green, puhescent and pale the upper surface olive green, completely gla­ below, thin chartaceous. Stipules 3-4 lines. brous, the lower surface appressed brownish Cymes tomentose, 1-lyz inch long, 3-5-flow­ villous on the principal veins, and closely white ered, the slender naked peduncle 6-15 lines, appressed pilose on the intervals; inflorescences the pedicels 1-lyz lines, bracteolate at the base. 3-6 mm long, axillary, spicate (or perhaps Calyx coriaceous, tomentose, 2 lines, shortly cymose); peduncle 2-4 mm long, densely ap­ 5-lobed, the tube continued beyond the ovary pressed brownish stiff villous; flowers 2-5, and in the fertile flowers constricted below the crowded; buds of perfect flowers ovoid sub­ throat. Disk plane and continuous without style globose; calyx in anthesis 3 mm long, densely in the sterile flowers. Style in the fertile flower appressed ascending brownish hirsutulous with­ short, with 3 obtuse stigmas." out, glabrous within; calyx lobes 1.2 mm long, EXPANDED DESCRIPTION FROM ALL SPECI­ deltoid ovate, thick; petals 0.4 mm long, 0.3 MENS EXAMINED: Shrub, decumbent or erect mm wide, spatulate, deeply hooded, enclosing and forming a rounded clump; stems 0.2-2 m the stamen; filament 0.4 mm long, lanceoloid, tall, to 1 cm in diameter, arching; branchlets attenuate; anther 0.25 mm long, obcuneate ob­ 2-4 mm in diameter, the older parts persistent long, retuse; disk with margin undulate and pilosulous; young branchlets ascending and 2-lobed opposite each calyx lobe; styles 2-3 half spreading dense pale to brownish pilosu­ and 0.6 mm long; immature capsule 4-5 mm lous; internodes 3-15 mm long, straight; nodes long, appressed pilosulous, orbicular in outline scarcely thickened; stipules 2-3 mm long, lance­ but sharply trigonous and with salient wings on linear, appressed white pilosulous, caducous, the angles, equally developed above and below. leaving a linear scar; petioles 8-35 mm long, HOLOTYPUS: Hawaiian Islands, Lanai, H. densely subappressed white pilose; blades 1.7­ Mmm & W. T. Brigham 344 (BISH). Isotypes 5.9 em long, 0.8-3.8 cm wide, chartaceous, (K, UC, us). broadly elliptic to elliptic, the base rounded, the apex shallowly emarginate (or in dwarfed DISCUSSION: Though the holotype and iso­ leaves minutely mucronate), the margin entire, types have several good branches in flower, the the upper surface olive green, glabrous, the inflorescences are small and young, and it has lower surface in youth densely appressed ascend­ not been possible to determine the exact kind of ing white pilosulous, but in age the hairs sexuality represented. spaced; inflorescences cymose, axillary, 15-45 520 PACIFIC SCIENCE, Vol. XXIII, October 1969

o 2",,,.· o Zmm ---~------" '--' '----' FIG. 6. Gouania Hillebfalldi Oliver, from Maui, lahainaluna. St. John ·25.609. a, Habit. X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal. X 30; e, stamen, X 30; t, capsule. X 4; g, seed, lateral view, X 5; h, seed and hilum, apical view. X 5.

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FIG. 7. Goual1ia Bishopii Hbd., from isotype, Maui, Lahaina, Bishop 43. a, Habit, Xl; b, bud of perfect flower, X 15; c, immature perfect flower, X 15; d, petal, X 30; e, stamen, X 30. 524 PACIFIC SCIENCE, Vol. XXIII, October 1969 pilosulous; internodes 4-25 mm long, straight; (Bishop Mus.) has a straight stem 26 em long, nodes scarcely swollen below the leaf scars; with 12 principal leaves. None of these have stipules "less than Yz inch [long], broad and tendrils. There are 9 axillary flowering branch­ obtuse," caducous, leaving a narrow lunate scar; lets and most of them with reduced leaves on petioles 5-15 mm long, terete, densely ascend­ the peduncle, but none bear tendrils. This ing rusty pilosulous; blades 4-8 em long, collection is indubitably an isotype, and there 2.8-4.8 em wide, elliptic to ovate, firm charta­ is no hint of a confusion. It seems that the ceous, the base shallowly subcordate, the apex species is a climbing shrub, with tendrils, but obtuse or subobtuse, the margins with larger that the tendrils are few, and when produced, hemispheric teeth alternating with smaller are only on the lateral flowering branchlets. dentate ones, all with salient blunt glandular Hillebrand dedicated the species to Edward tips, the upper surface dark green and moder­ Fletcher Bishop (1853-1875) of Lahaina, ately subappressed white pilosulous above, al­ Maui who had cooperated with him in the col­ most equally so on veins and intervals, the lower lection and study of the Hawaiian flora. surface pale green and with more dense similar pubescence; inflorescences terminal, on short 8. Gouania Lydgatei sp. nov. (Fig. 8, 17b) axillary branches with 1-2 leaves and, fide Hille­ G. Hillebrandi Oliver var. ~ Hillebrand, Fl. brand, "a stiff revolute tendril in their axils," Haw. Is. 83, 1888. the floriferous spicate rhachis 8-19 mm long, densely spreading rusty pilosulous, and bearing DIAGNOSIS HOLOTYPI: Frutex, ramulis 2-3 numerous crowded flowers; bracts 2.5-3 mm mm diametro densiter molliter subdivergente long, lance-ovate, with similar pubescence; buds pallidi-pilosis, internodis 2-10 mm longis rectis, 1.5-2 mm long, oblate globose, immature, ob­ nodis incrassatis, stipulis 2-3 mm longis anguste scured by the dense rusty pilosity of the calyx; lanceolatis adpresse albi-villosis caducis, petiolis calyx lobes 1.1 mm long, ovate concave, acute, 14-25 mm longis teretibus gracilibus densiter thick and coriaceous, glabrous within; calyx subadpressi-pilosulis, laminis 3-6.5 em longis tube 1.5 mm long, funnelform; immature petals 2-3.5 em latis firme subcoriaceis latiter ellipticis 0.7 mm long, spatulate, obovate, concave and in basi rotundatis in apice rotundatis sed minime deeply cucullate, white, hooded, enclosing the subcucullatis et revolutis marginibus integris stamen; filaments 0.3 mm long, broad subulate; supra obscure viridibus glabris sed midnervo ad anthers 0.25 mm long, wider than long, the two basim minime pilosulis infra densiter adpresse sacs ellipsoid; disk margin convex, emarginate, albis sericeis pilosulis pilis adscendentibus et then V-cleft to each filament base; styles 3 and cum nervis lateralibus paralle1is, inflorescentia 0.2 mm long (immature), then united below 2-5 em longis cymosis, pedunculis 10-40 mm for 0.2 mm. "Capsule tomentose, 3 lines high longis axillaribus adscendentibus gracilibus et and 4 lines wide, 3-winged, with wings con­ adscendente subadpresse albi-pilosulis, floribus tinued to the base, finally separating into 3 4-9 congregatis, rhachidi 2-7 mm longo pilo­ cocci, each coccus suspended by two filaments of sulo, bracteis 2-3 mm longis lanceolatis adpresse the 6-partite axis. Seeds plano-convex, brown­ pilosulis, alabastris 1-1.5 mm longis subglo­ ish." bosis dense adpresse hirsutulis, lobis calycis in floribus bisexualibus 1.5 mm longis deltoideis HOLOTYPUS: Flora Hawaiensis, W. Maui, intra glabris extra adscendente adpresse hirsu­ back of Lahaina, E. Bishop 43 (B)! Isotype tulis, tubo 1.2 mm longo semigloboso, petalis (BISH) . 1 mm longis spatulatis cucullatis et staminem The accompanying photograph of the holo­ inclusis, filamentis 0.6 mm longis subulatis, typic sheet once in Berlin shows one good leafy antheris 0.3 mm longis et latioribus thecis ob­ branch and five small detached branchlets. Seven lati-subglobosis in basi divergentibus, margini­ tendrils are visible, all originating from the axil bus disci ad filamentas V-fidis et inter filamentas of the ultimate leaf on a short, lateral, flowering concavis, stylis 3 et 0.3 mm longis in tertia in­ branchlet. None are produced from the principal fera connatis, capsulis 7-11 mm longis 7-9 mm leafy nodes of the main stem. The isotype diametro semiorbicularibus axili 6-7 mm longo Hawaiian Species of Gouania-ST. JOHN 525

FIG. 8. Gouania Lydgatei St. john, from holotype, Maui, Kula, Lydgate 134. a, Habit, X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; f, capsule, X 5; g, seed, lateral view, X 5; h, seed and hilum, apical view, X 5. 526 PACIFIC SCIENCE, Vol. XXIII, October 1969 corpore 3-laterato et 3-alato alis adscendente as the axis, the wing tapering downward to the arcuatis lanceolatis obtusis cuneatis dimidiis tam narrow base, the surface closely appressed white longis quam axili densiter adpresse albi-pilosulis, pilosulous; seeds 3.3-4 rom long, 1.5-1.6 mm seminibus 3.3-4 rom longis 1.5-1.6 mm latis wide, 0.5 mm thick, elliptic oblong, compressed, 0.5 mm crassis elliptici-oblongis compressis the longitudinal axis curved, the testa hard subcurvatis brunneis rigidis laevibus lucidis. smooth shining brown.

DIAGNOSIS OF HOLOTYPE: Shrub; branchlets HOLOTYPUS: Hawaiian Islands, E. Maui, 2-3 mm in diameter, densely soft half spreading lower part of Kula, J. M. Lydgate 134, ex Herb. pale pilose; internodes 2-10 mm long, straight; Hillebrand and Lydgate (BISH). Isotypes, Kula, nodes swollen around the leaf scars; stipules E. Maui, Herb. Hillebrand (BISH); and (B), 2-3 mm long, narrowly lanceolate, appressed see photo; and received July 1865, Dr. Hille­ white villous, caducous, leaving a short elliptic brand 125 (K)! scar; petioles 14-25 mm long, terete, slender, densely half appressed pilosulous; blades 3-6.5 DISCUSSION: The specimen in Bishop Museum cm long, 2-3.5 cm wide, firm subcoriaceous, labeled as an isotype of G. Hillebrandi consists broadly elliptic, the base rounded, the apex of a single detached capsule. From its morphol­ rounded except for the slight cucullate revolute ogy, it is clearly Hillebrand's var. ~, that is, the tip, the margins entire, the upper surface dark plant here described as G. Lydgatei. green, glabrous except that the midrib is slightly The well-developed flowers examined were pilosulous just at the base, below densely ap­ all perfect or bisexual. The available lateral pressed white silky pilosulous, the hairs ascend­ flowers on the four sheets are numerous, but ing parallel to the lateral veins; inflorescence mostly are too small and young to be worth 2-5 cm long, cymose; peduncle 10-40 mm long, dissecting. The largest one of these was boiled axillary, ascending, slender, and subappressed and dissected, and though but half grown, it ascending white pilosulous; flowers 4-9, seemed to be perfect also. In conclusion, the crowded; rhachis 2-7 mm long, similarly pilo­ mature flowers seen are perfect. Whether or not sulous; bracts 2-3 mm long, lanceolate, ap­ there are any imperfect flowers has not been pressed pilosulous; buds 1-1.5 mm long, sub­ determined. globose, obscured by the dense appressed The new epithet is chosen to honor the hirsutulosity; perfect flowers having the calyx Reverend John Mortimer Lydgate (1854-1922) lobes 1.5 rom long, deltoid, glabrous within, early assistant and long a collaborator with W. ascending appressed hirsutulous without; calyx Hillebrand in collecting and studying the flora tube 1.2 rom long, semiglobose; petals 1 mm of Kauai, Oahu, and Maui. long, spatulate, hooded, enclosing the stamen; filament 0.6 mm long, subulate; anthers 0.3 9. Gouania thinophila sp. nov. (Fig. 9a-f) mm long, wider than long, the two sacs oblate DIAGNOSIS HOLOTYPI: Frutex 13-19 cm altus, subglobose, divergent at base; disk margin V­ radici verticali elongati robusti, radicibus super­ cleft to each filament base, the rest of the mar­ ficialibus paucis, caule erecto unico vel cum gins shallow concave; styles 3 and 0.3 mm long, ramis paucis in basi 2.5-3 rom diametro teretibus united for lower ~; capsules 7-11 rom long, sublaevibus sed cortice brunneo cum squamis 7-9 mm in diameter, in outline semiorbicular, latis, ramulis foliosis 1.5 mm diametro dense the center axis 6-7 rom long, the body 3-sided subdivergenti-pilosis, internodis 1-6 rom longis and strongly 3-winged, the wings ascending rectis, nodis incrassatis et sub cicatrici projecten­ arcuate, lanceolate, obtuse, about half as long tibus, stipulis 2-3 mm longis caducis linearibus

FIG. 9. GOlta11ia thillophi/a St. John, from holotype, Maui, U.S. Exploring Expedition. a, Habit, XI; b, bud of perfect flower, X 15; C, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; f, seed, lateral view, X 5. Gouallia Hillebralldi Oliver, from Maui, Lahainaluna, St. John 26,719. g, Bud of staminate flower, X 15; h, staminate flower, X 15; i, petal, X 30; j, stamen, X 30. Hawaiian Species of GOllailia-ST. JOHN 527

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0...., I-'rn m 0..... ,:..,.",'" 528 PACIFIC SCIENCE, Vol. XXIII, October 1969 dense adpresse pilosis, petiolis 7-14 rnm longis lobes 1.8-2 mm long, ovate, subacute, thick and gracilibus et dense subadpresse fusci-pilosis, coriaceous, within glabrous and with a triangu­ laminis 1.4-3.5 cm longis 0.8-2.2 cm latis sub­ lar thickening running down 2/5 way from the coriaceis ellipticis planis integris supra obscure tip; petals 1 mm long, 0.4 rnm wide, white, lutei-viridibus glabris infra pallidioribus et ad­ spatulate, concave and deeply cucullate hooded, presse pilosulis in apice breve umbonato, in­ enclosing the stamen; filaments 0.7 mm long, florescentiis multis 7-15 rnm longis cymosis subulate; anthers 0.3 rnm long, suborbicular, densiter subadpresse albi-pilosulis 2-5-floriferis 2-lobed; disk concave, the margin notched to eis congregatis, floribus licet polygamis, floribus each filament base and retuse between each terminalibus perfectis et cum calycibus in flore pair; the three styles 0.5 rnm long, united below, 2.7-3 mm longis densiter subadpresse adscen­ 3-parted half way down; stigma apical discoid; dente albi-pilosulis tubo 1.3-1.5 mm longo immature capsules 6 mm long, obovoid sub­ obovoidei-hemispherico, 5 lobis 1.8-2 rnm longis orbicular, densely appressed white puberulent, ovatis subacutis crassis coriaceis intra glabris et with 3 lateral wings 1.5-2 mm wide, these parte 2/5 supera incrassata, petalis 1 rnm longis broadest near the apex, narrowing toward "the 0.4 mm latis albis spatulatis concavis et profunde base; lateral flower buds very immature, too cucullatis et staminem inclusis, filamentis 0.7 young to reveal whether perfect or staminate. mm longis subulatis; antheris 0.3 rnm longis suborbicularibus bilobatis, disco concavo mar­ HOLOTYPUS: Sandwich Islands [Hawaiian gine ad filamentas indentato et inter eas retuso, Is.]; Maui, sand hills, U.S. South Pacific Ex­ stylis 3 et 0.5 rnm longis parte Vz infera con­ ploring Expedition, 1838-42 (GH) . Isotype nata parte supera trilobata, stigmate apicali with only the locality, Sandwich Islands (us)! discoideo, floribus mascuIis ignotis; capsulis im­ On the sheets of this species in both the maturis 6 mm longis obovoidei-suborbicularibus herbaria there is another different species. In densiter adpresse albi-puberulentis 3-alatis, eis the Gray Herbarium, this has a separate label, 1.5-2 rnm latis ad apicem latioribus. Gouania orbicularis, Sandwich Islands. These extraneous specimens are here redetermined as DIAGNOSIS OF HOLOTYPE: Shrub 13-19 em G. Meymi Steud. tall; tap root elongate, strong; surface roots few; The new epithet is formed from the Greek, stem erect, simple or few-branched, at base this, sand hills, and phileo, to love, and is given 2.5-3 mm in diameter, terete, rather smooth, with reference to the habitat of the plant, on but checking into broad flakes, brown; leafy sand dunes. branches 1.5 mm in diameter, obscured by the dense, half spreading pilosity; internodes 1-6 10. Gouania pilata sp. nov. (Fig. 10) mm long, straight; nodes enlarged by a bracket­ DIAGNOSIS HOLOTYPI: Licet frutex, ramulis like swelling at the leaf scar; stipules 2-3 mm 2-2.3 rnm diametro densiter subadpresse albi­ long, linear, densely appressed pilose, caducous; pilosis et pilis persistentibus, internodis 12-38 petioles 7-14 mm long, slender, covered by a mm longis rectis, nodis subincrassatis, stipulis dense subappressed tawny pilosity; blades 1.4­ 4-8 mm longis caducis lancei-linearibus densiter 3.5 cm long, 0.8-2.2 cm wide, subcoriaceous, adpresse pilosis, petiolis 15-30 rnm longis elliptic, short umbonate, flat, entire, the upper teretibus gracilibus densiter adpresse albi-pilosis, surface dull yellowish green, glabrous, the lower laminis ramulorum principalium 5-6.5 cm longis surface slightly paler green, appressed white 1.6-2.5 em latis subtiliter chartaceis lanceolatis pilosulous, but the blade surface visible between vel anguste lanceolatis in basi rotundatis in apice the hairs; inflorescences numerous, 7-15 rnm anguste obtusis marginibus integris supra ob­ long, cymose, densely subappressed whitish scure viridibus omnino glabris infra ab initio pilosulous; flowers 2-5, crowded, apparently densiter adpresse adscendente albi-pilosis et in polygamous, the apical perfect flowers with midnervo et nervis principalibus cum pilis calyx in anthesis 2.7-3 rnm long, densely sub­ majoribus sed in foliis vetustioribus infra cum appressed ascending whitish pilosulous, the tube pilis separatis, inflorescentiis 22-35 mm longis 1.3-1.5 mm long, ovoid-hemispheric, the .'5 racemosis densiter subadpresse adscendente albi· Hawaiian Species of Gottania-ST. JOHN 529

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FIG. 10. Gouania pilata St. John, from holotype, Maui, Herb. Hillebrand. a, Habit, X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30. October 1969 530 PACIFIC SCIENCE, Vol. XXIII, with 5 broad pilosulis, pedunculo 13-30 mm longo gracili, mm long, free for upper half; disk pedicellis 1-2 mm longis, alabastris 1.3 mm retuse lobes. oblate ovoidei-subglobosis densiter ad­ longis HOLOTYPUS: Hawaiian Islands, Maui, ex lobis calycorum 1.3 mm presse albi-pilosulis, Herb. Hillebrand (BISH). There was also an intra glabris et midnervo longis ovati-ellipticis isotype (B)! Also W. & E. Maui, Hillebrand apiceque triangulari incrassatis, marginibus (GH). petalis 0.7 mm longis 0.3 mm latis obovate spatulatis marginibus apiceque involutis et DISCUSSION: This is a segregate from G. staminem inclusis, filamentis 0.5 mm longis Hillebrandi Oliver which was based upon two subulatis, antheris 0.25 mm longis thecis el­ Maui collections. The one from Lahaina, with liptici-oblongis in basi valde divergentibus, 3 broadly elliptic blades, is the lectotype of G. stylis 0.6 mm longis parte Yz supera libera, Hillebrandi. The one with lanceolate blades, disco cum lobis 5 latis retusis. labeled merely Maui (though Hillebrand cited it as from Kula), a specimen without collector shrub; DIAGNOSIS OF HOLOTYPE: Evidently a and without precise locality, is the holotype of branchlets 2-2.3 mm in diameter, densely sub­ our new G. pilata. the hairs long per­ appressed white pilose, There is nothing to indicate whether this 12-38 mm long, straight; sistent; internodes specimen was collected by Hillebrand himself, beneath the leaf scar; nodes slightly enlarged or by one of his collaborators. stipules 4-8 mm long, lance-linear, concealed The new epithet is the Latin adjective pilatm, by the dense appressed pilosity, early caducous, armed with a javelin, and is given with ref­ leaving an orbicular scar; petioles 15-30 mm erence to the shape of the lanceolate stipules. long, terete, slender, concealed by the dense blades of appressed ascending white pilosity; 11. Gouania Remyi sp. nov. (Fig. 11) main branches 5-6.5 cm long, 1.6-2.5 cm wide, Licet frutex, ramulis thin chartaceous, lanceolate to narrowly lanceo­ DIAGNOSIS HOLOTYPI: subadpresse persis­ late, the base rounded, the apex narrowly 2-3 mm diametro densiter internodis 6-10 mm longis obtuse, the margins entire, the upper surface tente albi pilosis, stipulis 3--4 mm dark green, wholly glabrous, the under surface rectis, nodis subincrassatis, valde adpresse at first hidden by the dense white appressed longis caducis lineari-Ianceolatis mm longis teretibus ascending pilosity, the midrib and laterals more albi-pilosis, petiolis 14-17 albi-pilosis, laminis coarsely pilose, but the older leaves with the gracilibus dense subadpresse cm latis subcoria­ hairs spaced and the under surface easily 3.2-3.7 cm longis 1.5-1.8 in basi rotundatis visible; inflorescences 22-35 mm long, race­ ceis ellipticis subacuminatis supra obscure viridibus in mose, densely subappressed ascending white marginibus integris pilis ad lateres disponitis pilosulous; peduncle 13-30 mm long, slender; toto adpresse pilosis secundariis cum pilis laxi­ pedicels 1-2 mm long; buds of perfect flower midnervo et nervis 25-35 mm longis axil­ 1.3 mm long, oblate ovoid-subglobose, densely oribus, inflorescentiis densiter subadpresse albi­ appressed white pilosulous; calyx lobes 1.3 mm laribus cymosis pedunculo 10-18 mOl long, ovate-elliptic, within glabrous and the pilosulis 5-11-floriferis, 1-7 mm longis, alabastris 2 midrib, margins, and apical triangle thickened; longo, pedicellis dense adscendente petals 0.7 mm long, 0.3 mm wide, obovate mm longis late ellipsoideis calycis 1 mm longis ovati­ spatulate, the margins and apex inrolled and albi-pilosis, lobis coriaceis intra glabris et enclosing the stamen; filaments 0.5 mm long, deltoideis crassis apiceque triangulari in­ subulate; anthers 0.25 mm long, the cells elliptic marginibus midnervo pilosis, petalis 0.6 mOl oblong, widely diverging at base; 3 styles 0.6 crassatis extra dense

Remy 587. a, Habit, X 1; b, bud of perfect FIG. 11. Gouania Remyi St. John, from holotype, Kahoolawe, stamen, X 30; f, capsule, X 4; g, staminate flower, flower, X 15; C, perfect flower, X 15; d, petal, X 30; e, X 30; k, smaller petal, X 30. X 15; h, larger petal, X 30; i, fertile stamen, X 30; j, sterile stamen, Hawaiian Species of GOltania-ST. JOHN 531

o

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" 5mm '-'~~-----'--'--'I 532 PACIFIC SCIENCE, Vol. XXIII, October 1969 longis ellipticis marginibus apiceque involutis angular thickening ~ as long as the lobe; et staminem inclusis, filamentis 0.5 mm longis petals 0.5-0.8 mm long, white, saccate cucul­ subulatis, antheris 0.2 mm longis et thecis late and enclosing a stamen; stamens unequal, oblongi-ellipsoideis in basi divergentibus, stylis two of them with anthers 0.3 mm long on fila­ 3 et 0.4 mm longis cylindricis crassis paene ments 0.2 mm long, three of them 0.5 mm parte Yz supera libera, disco undulato 10-lobato, long, containing pollen, and on filaments 0.3 capsulis 8 mm longis 7 mm latis subglobosis mm long; the 3 styles 0.2 mm long, separate, cuneatis dense subadpresse albi-pilosis axili each bearing an apical red stigma; disk 1.6 mm 6-6.5 mm longo 3-lateratis 3-alatis, alis in basi in diameter, almost circular, but notched to the 1 mm latis in apice 2 mm latis et cum loba 1 base of each filament, and the intervals with 2 mm longa ovata adscendenti. rounded lobes; capsules 8 mm long, 7 mm wide, cuneate, subglobose, densely white subappressed DIAGNOSIS OF HOLOTYPE: Apparently a shrub; pilose, the central axis 6-6.5 mm long, 3-sided branchlets 2-3 mm in diameter, densely sub­ and with 3 lateral wings 1 mm or less wide at appressed persistent white pilose; internodes the bottom, 2 mm wide at the top, and pro­ 6-10 mm long, straight; nodes slightly en­ jecting upwards as ovate lobes y!; as long as the larged around the leaf scar; stipules 3-4 mm axis. long, linear lanceolate, concealed by their white appressed pilosity, caducous, leaving a discoid HOLOTYPUS: lIes Sandwich, Kahoolawe, scar; petioles 14-17 mm long, terete, slender, 1851-1855, J. Remy 587 (p, the left hand concealed by the subappressed white pilosity; specimen). blades 3.2-3.7 em long, 1.5-1.8 em wide, sub­ DISCUSSION: The holotype of G. Remyi is the coriaceous, elliptic, the apex sharply sub­ left hand specimen on the sheet of Remy 587 in acuminate, the base rounded, the margins entire, the Paris herbarium. The sheet bears two the upper surface dark green, completely gla­ branches of quite different looking specimens brous, the lower surface concealed by the dense of Gouania. The right hand one is now marked white appressed pilosity, the hairs directed 587a, and is here described as G. cucullata. laterally, the midrib and secondaries with looser This species and G. cucullata, both from similar pubescence; inflorescences 25-35 mm Kahoolawe, are lost species and certainly extinct. long, axillary, cymose, densely subappressed The story of the utilization of the island of white pilosulous, bearing 5-11 polygamous Kahoolawe is a sad one of destructive exploita­ flowers; peduncle 10-18 mm long; pedicels tion. The island is eleven by six miles, and is 1-7 mm long; terminal flowers perfect, their 1,491 feet high. Being of only moderate height buds 2 mm long, broadly ellipsoid, concealed and situated in the lee of the 10,025-foot by the ascending white pilosity; calyx lobes 1 Haleakala Volcano, Maui, it receives only a low mm long, ovate deltoid, thick coriaceous, within rainfall, from 10-20 inches (in 1911-1917) glabrous and with thickened margins, midrib or 23-27 inches a year (in 1936-1938) fide and triangular apex, without densely pilose; Stearns (1940:124), and was not or little in­ petals 0.6 mm long, elliptic, with inrolled habited. Under the Hawaiian native kings it was margins and apex, hooded over and enclosing used as a convict colony as early as 1830, and the stamen; filaments 0.5 mm long, subulate; this continued at least until 1852. In 1832 anthers 0.2 mm long, of 2 oblong-ellipsoid sacs, there were 80 inhabitants, but in 1841 there diverging at base; styles 3 and 0.4 mm long, were only 15 convicts and a few fishermen. cylindric, stout, parted nearly half way; disk These settlers introduced pigs and goats and undulately 10-lobed; lateral flowers functionally used them for food, but apparently both es­ staminate, the buds 2.2 mm long, almost con­ caped and persisted, as did dogs. ,cealed by the appressed ascending white pilosity, Early accounts of the island describe it as the tube 1.2 mm long, hemispheric, the 5 lobes forested on top. Of this tree growth only 1 mm long, ovate, acute, coriaceous, within Erythrina sandwicensis persisted, and specimens glabrous and the midrib raised, the margins of it were collected by Forbes in 1913 from much thickened and the acute tip with a tri- one of the few surviving individual trees Hawaiian Species of Gouania-ST. JOHN 533

(1913:4,6, when still called E. monosperma). ranch. In 1863 the Hawaiian king leased the He learned from previous visitors that they had island for $250 a year to Charles G. Hopkins, also seen on the island the tree Reynoldsia and the lease was later assigned to a succession sandwicensis, and the shrubs Dodonaea sand­ of others, Elisha H. Allen, R. von Tempsky, A. wicensis, Styphelia Tameiameiae, Myoporttm McPhee, H. H. Baldwin, and others. The first sandwicense, and Euphorbia multiformis. An­ sheep ranch there failed. Later lessees introduced other shrub, Neraudia kahoolawensis, described sheep, cattle, horses, and pigs, and particularly by Hillebrand in 1888, had become extinct by the sheep and the goats multiplied and ran 1913. Among these species, the only real trees wild. In 1887 there were 900 cattle, and are the Erythrina and the Reynoldsia, both of 12,000 sheep. This overstocked the arid island, which will grow to good size, even in arid and the edible vegetation was soon consumed. places, but there they occur as scattered in­ The sod and the soil cover were destroyed, dividuals and can scarcely be said to produce especially by the sheep by their close cropping much of a forest. At least it is of a wide open and by their trampling, exposing the fine-par­ savannah type. The other known native plants ticled soil, which on the broad, flattish top of which are shrubs would have made some thickets the island was from 4 to 10 feet deep. Through­ between the trees, and filling in would be the out nearly all of the year the Hawaiian Islands common "pili" grass, Heteropogon contorttts. receive a brisk north-northeast trade wind. This These shrubs were the N eraudia, Dodonaea, hits Puu Kukui, the mountain mass of west Styphelia, Myoporum, and Euphorbia, and to Maui. Further east it hits the 10,025-foot these we now add Gouania Remyi and G. cuettl­ Haleakala in east Maui and in general fails to lata. Several other shrubs in the vegetation of blow over it; instead it divides and passes the littoral zone are here omitted because they around either flank. There is a broad isthmus would not have formed part of the upland between the two mountain masses, and the wind savannah forest vegetation. There was one other blows through this funnel-like opening in the endemic species collected by Remy, Lipochaeta direction of Kahoolawe. A turbulence of the kahoolawensis Sherff, and an indigenous one, air here is due to the current skirting the flank L. connata (Gaud.) DC. These are doubtless of the mountain, being diverted and mixing gone, but another endemic one was collected by with the straight trade passing over the Bryan in 1931, L. Bryanii Sherff. Together, isthmus. Those winds which climb on the then, the known endemics of the now desolated mountain slope lose most of their moisture island Kahoolawe are five: Gouania cucullata, there. After overpassing Maui, they blow on G. Remyi, Neraudia kahoolawensis, Lip0 chaeta and reach Kahoolawe, 6% miles to leeward. Bryanii, and L. kahoolawensis. Seldom do they bring rain to that bit of land. Jules Remy was in the Hawaiian Islands Any strong wind picked up the exposed fine from 1851 to 1855, and once during that soil of the upland of Kahoolawe and carried period he visited Kahoolawe. This was just at it for miles out to sea as a cloud of red dust the end of the penal colony period, and prob­ (Bryan, 1933:53). In this way all the good ably the natural vegetation was but little dis­ soil of the uplands down to bed rock, was turbed. He collected twelve plant species on it, blown away and lost. The lower slopes of the including the native shrubs Santalum ellipticum, island, where not cliffs, are rocky slopes. The Hibiscus Brackenridgei, and the two endemic general absence of soil on these slopes is ex­ species of Gouania here described. The Santa­ plained by Stearns as due to wave erosion during lum might have grown all over, from the shore a period when the island was submerged to the to the volcanic crests, and a single bush of it present 800-foot level (1940:147). This process was later found at the southwest end of the ruined the pasture and has nearly destroyed the island by Stearns (1940:124). The Hibiscus native flora. probably grew in rocky gulches in the upland. In 1910 the Territory of Hawaii repossessed It is a stout shrub or small tree, and may have the island, as a forest reserve, but the Board of added to the forest cover. Forestry was not given the finances and re­ The island was subsequently exploited as a sources with which to carry out a reclamation. 534 PACIFIC SCIENCE, Vol. XXIII, October 1969

Efforts to exterminate the stock then reduced pilose; major blades of main shoot 3-4.1 C111 these feral animals to 150 sheep and 4,300 long, 1.5-2.2 cm wide, thin subcoriaceous, goats. In 1939 the animal population was elliptic ovate, the base rounded, the apex acute estimated as 500 cattle, 200 wild sheep, 25 wild and apiculate, the margins entire, the upper goats, 17 horses, 3 mules, 500 wild turkeys, and surface dark green, completely glabrous, the plenty of rats, mice, and cats. The island is now lower surface with dense appressed whitish used by the U.S. Military as a bombing target, pilosity, the hairs directed laterally parallel to and is off limits to visitors. the lateral veins; inflorescences 5-10 mm long, The new epithet honors the French botanist, axillary, appressed pale pilosulous, spicate, Jules Remy (1826-1893), who lived on and ex­ bearing 3-5 flowers; peduncle 2-4 mm long; plored the Hawaiian Islands from 1851 to 1855. buds 1.5 mm long, subglobose, acute; calyx lobes His visit to Kahoolawe was fortunately just be­ 1.1 mm long, broadly ovate, acute, within fore the thousands of sheep and other grazing glabrous, the midrib, margins, and triangular animals were placed on the island. apex thickened, without appressed pilosulous; petals 0.6 mm long, 0.3 mm wide, obovate, the 12. Gouania cucullata sp. nov. (Fig. 12) margins and apex inrolled, cucullate, enclosing DIAGNOSIS HOLOTYPI: Licet frutex; ramulis the stamen; filament 0.3 mm long, subulate, 2.5-3 mm diametro densiter subadpresse pallide anther 0.3 mm long, ellipsoid-oblong, the sacs pilosis, internodis 3-7 mm longis, nodis paene only slightly divergent at base; disk only slightly incrassatis, stipulis 2-2.5 mm longis lanceolatis undulate; style 0.3 mm long, fully united; fruit obscuratis densiter adpresse pilosis caducis, peti­ unknown. olis 4-14 mm longis cernuis densiter subad­ HOLOTYPUS: Iles Sandwich, Kahoolawe, presse flavescentiter pilosis, laminis pricipalibus 1851-1855, J. Remy 587a (p, the right hand 3-4.1 cm longis 1.5-2.2 cm latis subtiliter sub­ specimen). coriaceis elliptici-ovatis in basi rotundatis in This species, endemic to Kahoolawe, is cer­ apice acutis et apiculatis marginibus integris tainly now extinct. For an account of the de­ supra obscure viridibus omnino glabris infra structive exploitation of this small island, see densiter adpresse albi-pilosis, pilis cum nervis the discussion under G. Remyi. lateralibus parallelis, inflorescentiis axillaribus The new epithet is the Latin adjective Cflcul­ 5-10 mm longis adpresse pallide pilosulis latus, hooded, and it is given with reference to spicatis et 3-5-floriferis, pedunculis 2-4 mm the hooded apex of the petals. longis, alabastris 1.5 mm longis subglobosis acutis, lobis calycis 1.1 mm Jongis latiter ovatis 13. Gouania hawaiiensis sp. nov. (Fig. 13) acutis intra glabris et in midnervo marginibus apice triangularique incrassatis extra adpresse DIAGNOSIS HOLOTYPI: lignosa cum pilosulis, petalis 0.6 mm longis 0.3 mm latis cirris, caulibus vetustioribus glabratis, cortice obovatis marginibus et apice involutis cucullatis griseo, ramulis densiter ferruginiter divergente et staminem inc1usis, filamentis 0.3 mm longis pilosulis, internodis 10-38 mm longis rectis, subulatis, antheris 0.3 mm longis ellipsoidei­ nodis incrassatis, stipulis caducis, petiolis 11-21 oblongis, saccis in basi subdivergentibus, disco mm longis, teretibus densiter divergente ferru­ subundulato, stylo non partitis, fructibus in­ giniter pilosulis supra canaliculatis, laminis 5.7­ 8.5 cm longis 4.2-6.5 cm latis crassis et firme cognitis. chartaceis latiter ellipticis (vel suborbicularibus) DIAGNOSIS OF HOLOTYPE: Evidently a shrub; in basi profunditer cordatis in apice obtusis vel branchlets 2.5-3 mm in diameter, densely sub­ breve subacutis marginibus breve rotunditer appressed pale pilose; internodes 3-7 mm long; dentatis supra obscure viridibus et persistente nodes slightly enlarged around the leaf scar; breviter subadpresse albi-pilosulis sed pilis non stipules 2-2.5 mm long, lanceolate, concealed congregatis nervis cum pilis simulantibus plur­ by the dense appressed pilosity, caducous, leav­ ibus densis infra albescenti-viridibus et spissiter ing an oval scar; petioles 4-14 mm long, pilosulis et in nervis optime pilosulis, cirris ex cernuous, densely subappressed pale yellowish apice ramulis lateralibus et 3-4 cm longis in- Hawaiian Species of GOltallia-ST. JOHN 535

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FIG. 12. Gouallia fUful/ala St. John, from holotype, Kahoolawe, Remy 587a. a, Habit, X 1; b, bud of perfect flower, X 15; f, perfect flower, X 15; d, petal, X 30; e, stamen, X 30. 536 PACIFIC SCIENCE, Vol. XXIII, October 1969

FIG. 13. Gouania hawaiiel/Sis St. John, from holotype, Hawaii Island, Keaa, Remy 588. a, Habit, X 1; b, leaf base, lower side, X 1. ··~·····' .. I ; 1~ ! i

o o.S".1I\ 0 , ! L..'_~_----' FIG. 14. Gouania sandwichiana St. John, from holotype, Hawaii Island, Remy 586. a, Habit, X 1; b, bud of perfect flower, X 15; c, perfect flower, X 15; d, petal, X 30; e, stamen, X 30; t, capsule, X 4; g, sterile flower, X 15; h, petal, X 30; i, stamen, X 30. 537 VI Ij.l 00

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...... \0 FIG. 15. Map of the distribution of the species of Gouania in the Hawaiian Islands. Numerals refer to numbered species described in text. 0\ \0 Hawaiian Species of Gottania-ST. JOHN 539 tegris intorquentibus densiter divergente fer­ and the hairs long persisting; internodes 10-38 rugini-pilosulis, spicis cum pedunculo 5 mm mm long, straight; nodes swollen below the longo, rhachidibus 20 mm longis ambis diver­ leaf scars; stipules caducous, leaving a narrow gente ferruginei-pilosulis, nodis 13, floribus et lunate scar; petioles 11-21 mm long, terete, fructibus perditis. channeled above, densely divergent rusty pilo­ sulous; blades 5.7-8.5 em long, 4.2-6.5 em DIAGNOSIS OF HOLOTYPE: Climbing cirrhifer­ wide, thick and firm chartaceous, broadly elliptic ous shrub; older stems glabrate, with gray bark; (or suborbicular), the base deeply cordate, the young stems densely divergent rusty pilosulous, apex obtuse or shortly subacute, the margins

FIG. 16a. Gouania Hillebrandi Oliver, holotype, Lahaina, Maui, herb. Hillebrand (Berlin). 540 PACIFIC SCIENCE, Vol. XXIII, October 1969 rounded short dentate, the surface above dark of Hawaii. With Remy's unique plant in mind, green and permanently subappressed short in 1965 William Meinecke searched in vain white pilosulous, but the hairs not crowded, the the rough lava flows and intervening thickets veins with similar but dense pubescence, below of that area near Waiohinu for this plant. the surface whitish green and shaggy pilosulous, The new epithet is a coined adjective, formed and all the veins outstandingly so; tendrils from of the place name, Hawaii, and the latin ad­ the tips of lateral shoots, 3---4 em long, un­ jectival place ending, -ensis. branched, coiling, densely ascending to diver­ gent rusty pilosulous; inflorescence spicate (or 14. Gouania sandwichiana sp. nov. (Fig. 14) perhaps racemose); peduncle 5 mm long; ~ rhachis 20 rom long, both divergent rusty G. orbicularis Walp. var. Mann, Proc. Am. pilosulous; nodes 13; flowers and fruit lost. Acad. Arts Sci. 7:162, 1867. DIAGNOSIS HOLOTYPI: licet frutex, ramis 2-3 HOLOTYPUS: Iles Sandwich, Hawaii 1, petit rom diametro obscure rubris et pilosis, ramulis bois au-dessus de Keaa, Kau, liane inconnue dense albi- vel brunnei-pilosis pilis in parte sub­ des indigenes, Juillet 1853, J. Remy 588 (p). divergentibus in parte subadpressis, internodis DISCUSSION: This seems closely related to G. 3-25 rom longis rectis, nodis subincrassatis, vitifolia Gray, of the Waianae Mountains, Oahu, stipulis 2-3 rom longis caducis anguste lanceo­ a species which has the blades 4-12 cm long, latis et densiter adpresse pilosis, petiolis 14-24 3--9 em wide, membranous, broadly cordate, mm longis teretibus gracilibus subadpressiter crenate, the upper half mostly narrowing to the albi- vel brunnei-pilosulis, laminis 2.6-5.1 cm tip; petioles subglabrate. G. hawaiiensis has the longis 1-2.7 em latis subtiliter chartaceis ellip­ blades 5.7-8.5 em long, 4.2-6.5 cm wide, thick ticis in basi rotundatis in apice rotundatis vel chartaceous, broadly elliptic (or suborbicular), minime emarginatis marginibus integris supra rounded dentate, the upper half rounding to olivaceis glabris infra palliditer viridibus et the tip; petioles with persistent shaggy pilosity. adpresse adscendente breve pilosis, inflorescentiis It is unfortunate that Remy's specimen shows 25-55 rom longis cymosis axillaribus, pedunculo only vegetative parts and a bare inflorescence 20---45 mm longo filiformi sparse subdivergente from which the flowers and fruit have fallen. pilosulo 5-13-florifero, floribus polygamis ab The land section Keaa is a small area in the dry inito congregatis tum separatis, pedicellis 1-5 lowlands near the southern end of the island mm longis dense semiadpresse pilosulis, ala-

FIG. I6b. Gouania Hillebrandi Oliver, at lahainaluna, Maui, St. John 26,723. Hawaiian Species of GOltallia-ST. JOHN 541 bastris ovoideis, floribus terminalibus perfectis obovatis albis marginibus apiceque involutis et cum calycibus in flore 2-2.3 mm longis densiter cucullatis et staminem inclusis, filamentis 0.8 adpresse adscendente brunnei-pilosulis lobis 1.3 mm longis crassis subulatis, antheris 0.4 mm mm longis ovatis acutus intra glabris et mid­ longis thecis binis oblongi-ellipsoideis in basi nervo marginibus apiceque triangulari incras­ divergentibus, disco cum 10 lobis latis rotun­ satis, petalis 1 mm longis 0.5 mm latis spatulati- datis, stylis 3 et 0.2 mm longis in toto connatis,

FIG. 17a. Gouallia BishoPii Hbd., holotype, lahaina, Maui, Bishop 43 (Berlin). 542 PACIFIC SCIENCE, Vol. XXIII, October 1969 capsulis 7-8 mm longis 6-7 mm latis sub­ nodes slightly enlarged beneath the leaf scars; orbicularibus in apice et basi emarginatis 3­ stipules 2-3 mm long, narrowly lanceolate, lobatis 3-alatis adpresse remote albi-puberulentis concealed by the appressed pilosity, caducous, alis 1.5-2.5 mm latis subsymmetricis. leaving a roundish scar; petioles 14-24 mm long, terete, slender, subappressed white to DIAGNOSIS OF HOLOTYPE: Evidently a shrub; brownish pilosulous; blades 2.6-5.1 em long, branchlets 2-3 mm in diameter, the older ones 1-2.7 em wide, thin chartaceous, elliptic, the dark reddish and with persistent pilosity; young base rounded, the apex rounded or minutely branchlets with abundant pilosity white to emarginate, the margins entire, the upper brownish, partly half spreading, partly sub­ surface olive green, glabrous, the lower surface appressed; internodes 3-25 mm long, straight; pale green and with dark reticulate Yenation,

FIG. 17b. Gouania Lydgatei St. John, holotype, Kula, Maui, Lydgate 134 (Berlin). Hawaiian Species of GOlfallia-ST. JOHN 543 and the surface with spaced, very short, ap­ CARLQUIST, SHERWIN. 1965. Island life. Garden pressed ascending white pilosity; inflorescences City, New York. 451 pp., illustrated. 25-55 mm long, cymose, axillary; peduncle ENGLER, A., and PRANTL, K. 1896. Die natiir­ 20-45 mm long, filiform, sparsely half spread­ lichen Pflanzenfamilien, 3(5): Rhamnaceae, ing pilosulous; cyme 5-13-flowered, the flowers by A. Weberbauer, pp. 393-427. Leipzig. polygamous, at first crowded, but in young fruit --- 1953. Die natiirlichen Pflanzenfamilien, separated; pedicels 1-5 mm long, densely semi­ 2nd ed. (H. Harms, editor). 20d: Rham­ appressed pilosulous; buds ovoid; terminal naceae by K. Suessenguth, pp. 1-173. Berlin~ perfect flowers with the calyx in anthesis 2-2.3 FORBES, CHARLES N. 1913. Notes on the flora mm long, densely appressed ascending brownish of Kahoolawe and Molokini. B. P. Bishop pilosulous; calyx lobes 1.3 mm long, ovate, Museum, Occasional Papers, vol. 5, no. 3, pp. acute, within glabrous, and the midrib, margins, 1-10, figs. 1-6. and apical triangle thickened; petals 1 mm long, GRAY, ASA. 1854. United States Exploring Ex. 0.5 mm wide, spatulate obovate, white, the pedition during the years 1838-1842, vol. margins and apex inrolled, hooded and enclos­ 15, pt. 1: Botany, pp. 1-777. ing the stamen; filament 0.8 mm long, stout HILLEBRAND, WILLIAM. 1888. Flora of the subulate; anthers 0.4 mm long, the 2 oblong­ Hawaiian Islands. London, New York, and ellipsoid sacs divergent at base; disk with 2 Heidelberg. 673 pp. broad rounded lobes opposite each calyx lobe; 3 styles 0.2 mm long, fully united; lateral LEVEILLE, H. 1912. Decades plantarum no­ flowers evidently sterile, the calyx 3 mm long, varum XC-XCII. 1. Plantae sandwicenses. Fedde Repertorium Novarum Specierum densely appressed ascending brownish pilo­ Regni Vegetabilis, vol. 11, pp. 63-67. sulous without, the tube 2 mm long, funnel­ form, the 5 lobes 1 mm long, valvate, broadly MEYEN, F. J. F. 1835. Reise urn die Erde ovate, acute, within glabrous, fleshy and thick­ ausgefiihrt auf dem Koniglich Preussischen ened at the margins, up the narrow midrib, and Seehandlungs-Schiffe Prinzess Louise, Com­ at the triangular apex; petals 0.7 mm long, andort von Capitain Wendt, 1830-1832. obovate saccate, deeply cucullate and enclosing Vol. 2. Berlin. 413 pp. a stamen; filaments 0.5 mm long, acicular; ROCK, J. F. 1914. Revisio plantarum Hawai­ anthers 0.3 mm long, cordate suborbicular; iensium a Leveille descriptarum. Fedde disk 1. 7 mm in diameter, with a broad, emar­ Repertorium Novarum Specierum Regni ginate lobe opposite each calyx lobe; the 2 Vegetabilis, vol. 13, pp. 352-361. styles 0.2 mm long, distinct, subcapitate; ovary ST. JOHN, HAROLD. 1952. Monograph of the evidently sterile; capsules of perfect flowers genus Isodendrioll (Violaceae), Hawaiian 7-8 mm long 6-7 mm wide, suborbicular, plant studies 21. Pacific Science, vol. 6, pp. emarginate at both ends, appressed white 213-255, figs. 1-15. spaced puberulent throughout, 3-lobed, 3­ STEARNS, HAROLD T. 1940. Geological and winged, the wings 1.5-2.5 mm wide, of nearly ground-water resources of the islands of Lanai equal width from base to tip. and Kahoolawe, Hawaii. Hawaii Territory, Division of Hydrography, Bulletin 6, 173 pp., HOLOTYPUS: Iles Sandwich, Hawaii 1., 1851­ 34 figs., 1 map. 1855, Remy 586 (p). Isotype (GH)! J. STEUDEL, ERNEsTo THEOPH. 1840. Nomen. DISCUSSION: The new epithet is formed of clator botanicus, 2nd ed. Vol. 1. Stuttgart and the geographic name Sandwich, and the Latin Tiibingen. 852 pp. adjectival suffix -iamfS. WALPERS, G. 1843. F. J. F. Meyenii observa· tiones botanicas in itinere circum terram in­ LITERATURE CITED stitutas. Nova Acta Academiae Caesareae BRYAN, EDWIN H., JR. 1933. Hawaiian nature Leopoldino-Carolinae Naturae Curiosorum, notes. Honolulu. 285 pp., illustrated. vol. 19, suppl. 1, pp. 1-512, figs. I-XIII.