MANUAL OF

THE FLOWERING PLANTS

OF HAWAI'I

Bishop Museum Special Publication 97

University of Hawai'i Press

Bishop Museum Press

page 241

11. ASTERACEAE Sunflower family

Herbs or sometimes shrubs, rarely small to medium-sized trees or lianas, sap watery of milky. Leaves simple or compound, alternate, opposite, or rarely whorled, stipules absent. Flowers (florets) borne in 1 to numerous dense heads with 1 to numerous sessile flowers on a common receptacle, heads subtended by an involucre of 1 to several series of bracts, sometimes clustered into secondary heads, often with a secondary involucre; receptacle flat, convex, concave to conical or cylindrical, the surface smooth, rough, or pitted, naked or chaffy (at least some florets subtended by a bract), occasionally densely bristly, the bristles usually not strongly associated with individual florets; florets perfect or unisexual and the marginal florets pistillate or neutral and the central ones perfect or functionally staminate; calyx absent or strongly modified in various ways and referred to as a pappus, inserted at apex of ovary, consisting of 1 to several or numerous scales, awns, or capillary or plumose bristles, occasionally the scales connate to form a low crown, sometimes consisting of more than 1 series of pappus type; corolla (3)4-5-lobed, connate most of its length, usually of 1 or 3 types: (a) usually perfect, actinomorphic, (3)4-5-lobed, the lobes equal and much shorter than the conspicuous tube, called disk florets, (b) zygomorphic, with a short tube and a flat, strap-like, usually 3-toothed limb (5-toothed in subfamily Cichorioideae), termed ray florets, that are often pistillate or neutral and lack stamens (always perfect in subfamily Cichorioideae), sometimes the strap-shaped limb is very short, 0.5-2 mm long, and the heads appear disciform, and (c) zygomorphic and bilabiate, the outer limb usually the largest, or actinomorphic, tubular as in type a, but the corolla lobes long and slender, sometimes unequaly split; ray florets, corolla type b, are often the peripheral in the head or make up the entire head; disk florets, corolla type a, are central in the head or constitute the entire head; stamens (4)5, alternate with the corolla lobes; filaments distinct or rarely connate, inserted on lower part of the corolla tube; anthers flat, elongate dithecal, usually with short apical appendage, obtuse to sagittate at base, connate into an anther tube, opening by longitudinal slits, releasing pollen into the tube which is then pushed out by the growth of the style, in Ambrosia and Xanthium the anthers distinct or loosely coherent; ovary inferior, 2-carpellate, 1-celled, ovule 1, basally attached and anatropous; style 1, 2-branched above, often not separating in functionally staminate

page 242

florets, stigmatic in lines or across the entire surface or upper portion of style, the branches often continuing above the stigmatic area to form sterile, often pubescent appendages. Fruit an achene, cylindrical, prismatic, conical, or compressed, often with the papus persistent. Seed 1, endosperm absent or essentially so. [Cronquist, 1980; Fosbert & Sachet, 1980a; Woodson, Schery & collabortors, 1975].

The family comprising 1,100 genera and some 20,000 species, cosmopolitan in distribution but most abundant in temperate and subtropical regions, Asteraceae are one of the most successful families of flowering plants. Few are forest trees or aquatics, but otherwise they are highly diversified in their habitats, life-forms, pollination mechanisms, and dispersal patterns. The family may be organized into about 13 tribes. One of these, Lactuceae, is often considered to be in a distinct subfamily, Cichorioideae; the remainder, which are largely overlapping, are in Asteroideae.

Represented in Hawai'i by 76 genera and 181 species distributed in 11 of the 13 tribes. The 91 native species are distributed in 12 genera. All of these except Adenostemma lavenia are endemic. The genera Argyroxiphium, Dubautia, Hesperomannia, Lipochaeta, Remya, and Wilkesia are endemic to Hawai'i, while Lagenifera and Tetramolopium, although not endemic, are represented by endemic species, and Artemisia, Bidens, and Gnaphalium have both naturalized and native species. Ninety species in 67 genera have become naturalized in Hawai'i; many are common weeds, but few have become serious problems.

The monotypic genus Zemisne Degener & Sherff was described (Sherff, 1935a; Degener, 1939a) from a single A. Menzies collection presumed to be from Hawai'i (Z. menziesii Degener & Sherff). The collection was evidently mislabeled and was actually from the Galápagos Islands (Sherff, 1947b; St. John, 1973a). It was renamed Scalesia menziesii (Degener & Sherff) Degener & Sherff in Sherff (1947b), but according to St. John (1973a), this is a later synonym of S. aspera Anderss. Another species, Dichrocephala integrifolia (L. fil.) Kuntze [Cotula bicolor Parkinson; Dichrocephala latifolia (Desf.) DC] has been reported from Hawai'i (Seemann, 1866a, p. 144; Fosberg & Sachet, 1981; and on a single David Nelson collection). This report probably represents a misidentification of Adenostemma lavenia. Boltonia indica (L.) Benth. was reported as a garden escape in Kāne'ohe, O'ahu, by Nagata (1987), but it does not appear to be fully naturalized. Hillebrand (1888) treated Cosmos caudatus Kunth as an ornamental species naturalized at least on Hawai'i. This mexican species is an annual with white to pink rays. We have not seen any Hawaiian collections, nor does it currently appear to be in cultivation. St. John (1973a) reports 2 other species as adventive. Rudbeckia laciniata L. cv. Hortensia, golden flow, and Tanacetum vulgare L., common tansy, both have occasionally been cultivated in Hawai'i; however there is no evidence that they have ever escaped cultivation. Neal (1965) also reports T. vulgare as occasionally escaping cultivation.

page 420

22. CAMPANULACEAE Bellflower family

Contributed by Thomas G. Lammers

Herbs, shrubs, or trees, branched or unbranched, unarmed or rarely muricate, usually with milky sap. Leaves simple, alternate, entire or toothed, rarely pinnately lobed or parted or nearly compound, stipules absent. Flowers perfect or rarely unisexual, sometimes resupinate, usually protandrous, rarely cleistogamous, solitary in the axils of cauline leaves or arranged in terminal or axillary racemes, spikes, or cymes; sepals connate, tube adnate to ovary, 5-lobed; corolla actinomorphic or, if zygomorphic, then often laterally fenestrate or dorsally cleft, connate with 5 valvate lobes; stamens 5, alternate with corolla lobes; anthers dithecal, opening by longitudinal slits, coherent but separating after anthesis or connate and forming a tube into which pollen is shed; filaments distinct or connate above, attached to the epigynous nectary disk or to base of corolla, rarely adnate to corolla tube; ovary inferior sometimes only partly so, rarely almost superior, 2-5-carpellate, with as many cells as carpels, placentation axile, or when 1-celled with 2 parietal placentas, often crowned with an annular nectary disk; stigmas wet or dry, 2-5-lobed, appressed and nonreceptive as the style grows through the anther tube, pushing out the pollen, after which the stigmas spread apart and become receptive. Fruit a berry or a loculicidal, poricidal, or rarely circumscissile capsule. Seeds numerous, small, with a straight, short to spathulate dicotyledonous embryo embedded in oily endosperm. [Baillon, 1885; Bentham, 1876b; A. P. de Candolle, 1839a; Carlquist, 1956, 1962a, 1969a; Carr, 1978; Cory, 1984; Don, 1834a; Drake del Castillo, 1890b, 1892; Ehrendorfer, 1979; Erdtman, 1952; Fosberg, 1948a; Gray, 1861b; Hooker & Arnott, 1832; Lammers, 1988a, 1988b; Lammers & Freeman, 1986; Mann, 1867; Presl, 1836; Rock, 1913b, 1919a; Schönland, 1889, Selling, 1947; Swezey, 1954; Wimmer, 1943, 1953, 1968]

page 421

A family comprising 70 genera and 2,000 species, cosmopolitan in distribution. Two subfamilies are recognized: Campanuloideae characterized by actinomorphic, nonresupinate flowers, coherent anthers, and 3-5-celled ovatires; the Lobelioideae charactered by zygomorphic, resupinate flowers, connate anthers, and 2-celled ovaries. Represented in Hawai'i by 8 genera of Lobelioideae (6 endemic, one indigenous, one naturalized) comoprising 110 endemic woody species and 2 naturalized herbaceous species; and 2 naturalized genera of Campanuloideae, each represented by a single herbaceous species. In addition to the species included in the treatment, Lobelia tenuior R. Br., Platycodon grandiflorum (Jacq.) A. DC, and a few species of Campanula L. are occasionally cultivated in Hawai'i (Neal, 1965).

The endemic Lobelioideae are truly the "peculiar pride of our flora" (Hillebrand, 1888, p. xxiv), evoking considerable interest from amateur and professional botanists alike. This interest is due largely to their extreme morphological diversity and realization that all this diversity stems from no more than 5 original colonization events (Rock, 1919a; Lammers & Stein, in prep.), one for each of the following groups: (1) Brighamia; (2) Clermontia, Cyanea, Delissea, and Rollandia; (3) Lobelia sect. Galeatella; (4) Lobelia sect. Revolutella; and perhaps (5) Trematolobelia. Of special interest is the role that endemic nectar-feeding birds, the honeycreepers, may have played in their pollination, and consequently in the evolutionary development of this diversity (Lammers & Freeman, 1986). It is tragic that nearly 25% of the spcies have become extinct in the century since Hillebrand first called attention to these botanical marvels.

KEY TO THE GENERA OF CAMPANULACEAE

page 422

1. BRIGHAMIA A. Gray Ālula, hāhā

Unbranched stem succulents 1-5 m tall; stems stout, fleshy, thickened toward base, glabrous the wood soft. Leaves simple, arranged in a dense apical rosette, fleshy, obovate, margins entire or weakly and distantly toothed in the apical 1/2, apex subacute to obtuse, base cuneate, sessile to subsessile. Flowers fragrant, 3-8 in suberect to horizontal axillary racemes; hypanthium cylindrical, 10-ribbed; calyx lobes shorter than hypanthium; corolla white or yellow, salverform, the tube slender, straight, entire at anthesis, later dorsally cleft ca. 1/6 its length, with age up to ca. 1/3 its length, the lobes subequal, their margins finely erose; staminal column adnate to corolla tube below the middle, glabrous to pubescent, not exserted; all 5 anthers with apical tufts of white hairs; ovary inferior, 2-celled; style exserted from corolla; stigmas 2-lobed. Fruit a capsule, ellipsoid, compressed, at first fleshy, eventually drying, each cell dehiscent by 2 lateral longitudinal slits. Seeds numerous, pale, ovoid to ellipsoid, smooth to minutely papillose. [Forbes, 1913a, 1917b; M. A. Johnson, 1984, 1986; Mabberley, 1975a; Mann, 1869; Rauh, 1981; Rowley, 1983, 1987; St. John, 1958, 1969a; Stone, 1957]

This endemic genus of 2 species is certainly one of the most unusual members of the Hawaiian flora. The combination of stem succulent habit, axillary inflorescences, and salverform corolla is unique among the Lobelioideae, making it difficult to satisfactorily elucidate relationships with other genera. The name commemorates William Tufts Brigham (1841-1926), first director of the Bishop Museum, who botanized Hawai'i in 1864-1865 with Horace Mann, Jr. (Mann, 1867; Rose, 1980).

KEY TO THE SPECIES OF BRIGHAMIA

page 423

1. Brighamia insignis A. Gray

[B. citrina (C. Forbes & Lydgate) St. John; B. c. var. napaliensis St. John; B. insignis f. citrina C. Forbes & Lydgate]

(end, E*) 'Ōlulu, pū aupaka

Leaves 12-20 cm long, 6.5-11 cm wide. Peduncule plus rachis 4-12 cm long, pedicels 10-30 mm long; hypanthium 10-12 mm long, 4.5-7 mm wide; calyx lobes deltate to ovate, 02.-1 mm long; corolla yellow, the tube 7-14 cm long, 0.3-0.4 cm wide, the lobes elliptic, 1.3-2.2 cm long, 0.7-1.1 cm wide; anthers glabrous or pubescent along the connectives. Capsules 13-19 mm long, 9-13 mm wide, 4-5 mm thick. Seeds 0.8-1.2 mm long, conspicuously papillose. [2n=28*.] Occurring on sea cliffs, 0-400 m, Ka'ali Cliff, Ni'ihau, and the Nāpali Coast and Hā'upu Ridge, Kaua'i.

St. John (1969a) segregated the Kaua'i populations as Brighamia citrina on the basis of minor differences in the pubescence of the staminal column and the shape of the papillae of the seed. However, the Ni'ihau and Kaua'i populations are here judged conspecific on the basis of their common possession of yellow corollas, papillose seeds, pedicels 1-3 cm long, and deltate to ovate calyx lobes up to 1 mm long. Last observed on Ni'ihau in 1947 (St. John, 1969a).

2. Brighamia rockii St. John

[B. remyi St. John; B. rockii f. longiloba St. John]

(end, E*) Pua 'ala

Leaves 6-22 cm long, 5-15 cm wide. Peduncle plus rachis 3.5-7.5 cm long, pedicels 6-12 mm long; calyx lobes oblong to elliptic, 2.5-8 mm long; corolla white, the tube green to yellowish green, 8-13 cm long, 0.2-0.4 cm wide, the lobes elliptic, 1.7-3.7 cm long, 0.8-1.3 cm wide; anthers glabrous. Capsules 13-20 mm long, 7-10 mm wide, 3-4 mm thick. seeds smooth, 1.1-1.2 mm long. Occurring on sea cliffs, 0-470m, windward coast of Moloka'i from Kalaupapa to Hālawa. It is possible that this species formerly occurred on Maui and Lāna'i as well. — Plate 48.

St. John (1969a) based Brighamia remyi on a single specimen (Rémy 309bis, P) collected on Maui in the early 1850s. Although the specimen lacks capsules and seeds and the color of the corolla is uncertain, it is here assigned to B. rockii on the basis of its oblong-elliptic calyx lobes 4.5-8 mm long and its pedicels 6-12 mm long. Rock (1919a) observed an inaccessible Brighamia at Maunalei Valley, Lāna'i, which may also be referable to this species.

2. CLERMONTIA Gaud. Ōhā wai, 'ōhā, hāhā

[Delissea Gaud. sect. Clermontia (Gaud.) Baill.]

Branched shrubs or small trees, terrestrial or epiphytic. Leaves callose-toothed. Flowers 2-3(-10) in axillary, subumbellate racemes; hypanthium hemispherical, obovoid, obconical, or turbinate; calyx lobes either mimicking the corolla (as long as corolla or nearly so, of similar shape, texture, and color, connate, deciduous) or similar to those found in other genera (shorter than hypanthium to 1/2 as long as the corolla, triangular to oblong, firm, green, distinct or rarely connate at base, persistent); corolla zygomorphic, the tube sub-erect to arcuate, dorsally cleft to near base, the lobes erect, downcurved, spreading, or recurved; stamens connate; anthers with apical tufts of white hairs on the lower 2; ovary inferior, 2-celled; stigma 2-lobed. Fruit an orange or yellow berry. Seeds dark brown to black, smooth, shiny, 0.2-0.5 mm long. [Dadswell & Record, 1936; Endlicher, 1836; Forbes, 1912, 1920; Gaudichaud-Beaupré, 1829a, 1841-?1852; Gray, 1861b; Heller, 1897; Herbst, 1980; Hochreutiner, 1934; Lammers, Weller & Sakai, 1986; Léveillé, 1911b, 1913b; Mabberley, 1975a; Meyen, 1843; Nuttall, 1843; Rock, 1957, 1962a; St. John, 1939a, 1939b, 1969b, 1971a, 1976a, 1977c, 1978c, 1978d, 1979e, 1980d, 1980e, 1983b, 1985a, 1987a; Skottsberg, 1926, 1944a, 1953; Spieth, 1966; Swanholm, St. John & Scheuer, 1959, 1960; Wawra, 1873b, 1873c; Wimmer, 1929]

page 424

page 425

An endemic genus of 22 species, divided into 2 sections: Clermontioideae (Hillebr.) Rock with calyx of the type found in other lobelioid genera (the lobes are green, triangular to oblong, distinct or rarely connate at base, persistent, shorter than the hypanthium to 1/2 as long as the corolla); and Clermontia with the calyx lobes mimicking the corolla, becoming as long as the corolla or nearly so, and matching it in color, shape, dehiscence, and degree of connation. The latter type creates a unique "double-flowered" perianth, comprising an inner series (corolla) of 2 lateral and 3 ventral loes, and an alternating outer series (calyx) of one dorsal and 4 ventral loes. the dorsal lobe is typically distinct almost to the base while the remaining lobes are typically distinct 1/6-1/2 their length (i.e., the perianth lobes are 1/5 as long to as long as the perianth tube.)

Clermontia, Cyanea, Delissea, and Rollandia have in common a suite of features unique among Lobelioideae: woody habit, axillary inflorescenses, and baccate fruit. This suite of features, together with their common chromosome numer (2n = 28), suggests that these 4 endemic genera are derived from a single ancestral stock. The Andean genera Burmeistera Triana and Cetropogon K. Presl have generally been implicated as possible ancestors (Rock, 1919a; Stone, 1967b; Carlquiest, 1974). A reconsideration of the data, however, suggests that Clermontia and its allies are most closely related to certain Indo-Malayan species of Pratia Gaud. sect. Colensoa (J. D. Hook.) Hemsl. (Lammers & Stein, in prep.) The genus commemorates M. le Marquis de Clermont-Tonnerre, Minister of the French Navy at the time of the Freycinet expedition (1817-1820).

KEY TO THE SPECIES OF CLERMONTIA

page 426

1. Clermontia arborescens (H. Mann) Hillebr.

[Cyanea arborescens H. Mann; Clermontia arborescens subsp. waihiae (Wawra) Lammers; C. furcata F. Wimmer; C. mannii St. John; C. waikoluensis St. John; Delissea waihiae Wawra]

(end) 'Ōhā wai nui

Terrestrial or epiphytic trees or shrubs 1.5-8 m tall. Leaves oblong, elliptic, or oblanceolate, blades 10-23 cm long, 2-6.5 cm wide, upper surface dark green, glossy, glabrous, lower surface greenish white, glabrous or sparsely pubescent along the veins, margins callose-crenulate, apex obtuse, acute acuminate, or cuspidate, base cuneate to attenuate, petioles 1.5-8 cm long. Inflorescences 1-2-flowered, peduncles 4-18 mm long, pedicels 12-40 mm long; hypanthium hemispherical or obconical, 10-20 mm long, 10-20 mm wide; calyx lobes triangular, deltate, shallowly triangular, oblong, or ovate, 3-40 mm long, 3-10 mm wide, apex acute, obtuse, or emarginate and often apiculate; corolla greenish externally, greenish white or magenta within, 55-85 mm long, 8-18 mm wide, glabrous or rarely pubescent, the tube suberect to curved, the lobes thick, succulent, arcuate, the sinuses separating each lateral lobe from the 3 ventral lobes 3-5 times longer than the sinuses among the ventral lobes; staminal column magenta, included within corolla, glabrous; anthers dark purple, glabrous or rarely sparsely pubescent. Berries orange, subglobose to obovoid, 25-40 mm long, 25-40 mm wide. Occurring in mesic to wet forest, 520-1,825 m, on Moloka'i, Lāna'i, and Maui. — Plate 48.

page 427

Two subspecies may be distinguished: subsp. arborescens [incl. Clermontia furcata, C. mannii] with calyx lobes oblong, ovate, or triangular, 10-40 mm long, apex acute to obtuse (southern West Maui); and subsp. waihiae [incl. Clermontia waikoluensis] with calyx lobes shallowly triangular, 3-9 mm long, apex emarginate or obtuse (Moloka'i, Lāna'i, northern West Maui, and East Maui). Reports of Clermontia arborescens from Kaua'i (Wimmer, 1943, 1968; Rock, 1957) are based upon misidentified specimens of C. fauriei.

2. Clermontia calophylla F. Wimmer

[C. montis-loa Rock var. tenuifolia Skottsb.; C. parviflora Gaud. ex A. Gray var. grandis Rock; C. p. var. intermedia Skottsb.]

(end)

Terrestrial shrubs 1.5-5 m tall. Leaves broadly elliptic to elliptic, blades 12-19 cm long, 4-7 cm wide, upper surface green, glabrous, lower surface pale green, glabrous, or pubescent, margins callose-crenulate, apex acuminate, acute, or obtuse, base cuneate, petioles 3-7 cm long, glabrous or pubescent. Inflorescences 2(-4)-flowered, peduncles 10-17 mm long, pedicels 10-12 mm long; hypanthium obovoid, turbinate, or hemispherical, 10-17 mm long, 8-12 mm wide, glabrous or pubescent; perianth greenish externally, violet or dark purple within, 35-45 mm long, 7-10 mm wide, glabrous or pubescent, the tube suberect, the lobes recurved, about as long as the tube; staminal column violet or dark purple, exserted, glabrous; anthers dark purple, glabrous. Berries orange, subglobose, 15-30 mm long, 15-30 mm wide. [2n = 28*.] Occurring in wet forest, 885-1,460 m, Kohala Mountains, Hawai'i.

This species has generally been treated as a variety of Clermontia parviflora. While it resembles that species in its recurved perianth lobes, it is amply distinct with its larger, dark purple perianth, larger hypanthium, wider leaves, and larger berries.

3. Clermontia clermontioides (Gaud.) A. Heller

[Delissea clermontioides Gaud.; Clermontia clermontioides subsp. rockiana (F. Wimmer) Lammers; C. coerulea Hiller. C. c. subsp. brevidens (Skottsb.) St. John; C. c. subsp. b. f. flavescens (F. Wimmer) St. John; C. c. var. brevidens Skottsb; C. c. var. degeneri Skottsb.; C. c. f. flavescens F. Wimmer; C. c. var. greenwelliana F. Wimmer; C. c. var. parvifolia Rock; C. gaudichaudii Hillebr., nom. illeg.; C. hualalaiensis St. John; C. konaensis St. John; C. loyana Rock; C. rockiana F. Wimmer]

(end)

Terrestrial or epiphytic shrubs or trees 1.5-6.5 m tall. Leaves oblong, elliptic, or oblanceolate, blades 7-25 cm long, 2-5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent along the veins, margins callose-crenulate, apex acuminate, acute, obtuse, or cuspidate, base cuneate or attenuate, petioles 1-7 cm long. Inflorescences 1-2(-4)-flowered, glabrous or pubescent, peduncles 18-50 mm long, pedicels 5-40 mm long; hypanthium hemispherical to obconical, 10-15 mm long, 10-15 mm wide; calyx lobes narrowly triangular to triangular, 2-36 mm long, 0.5-10 mm wide, distinct or connate at base, apex acute or acuminate; corolla greenish, white, or purplish, 40-60 mm long, 6-12 mm wide, glabrous or pubescent, the tube suberect to curved, the lobes all of equal length, about as long as the tube, spreading; staminal column white or purplish, glabrous; anthers white or purplish, glabrous. Berries orange or yellow, subglobose, 20-36 mm long, 20-36 mm wide. [2n = 28*.] Occurring in mesic to wet forest, 670-1,825 m, Ka'ū and Kona districts, Hawai'i. — Plate 48.

Two subspecies may be distinguished: subsp. clermontioides [incl. Clermontia coerulea, C. c. var. brevidens, C. c. var. degeneri, C. c. f. flavescens, C. c. var. parvifolia, C. konaensis] with calyx lobes distinct, 2-10 mm long, 0.5-3 mm wide (Pāhala to Kealakekua, 670-1525m); and subsp. rockiana [incl. C. coerulea var. greenwelliana, C. hualalaiensis, C. loyana] with calyx lobes connate at base, 10-36 mm long, 3-10 mm wide (Keālia to Hualālai, 840-1825 m). Where the ranges overlap (Keālia to Hōnaunau), subsp. clermontioides tends to occur at lower elevations than subsp. rockiana This species has been known as Clermontia coerulea for many years. However, the name Clermontia clermontioides, long misapplied to the plants of Kaua'i and O'ahu now called C. fauriei, has priority (Lammers, 1988b).

page 428

4. Clermontia drepanomorpha Rock

(end)

Terrestrial or epiphytic trees 2-7 m tall. Leaves oblanceolate or elliptic, blades 8-22 cm long, 1.5-4.5 cm wide, upper surface green, glabrous, lower surface pale green or purplish, glabrous or sparsely pubescent along the midrib, margins callose-crenulate, apex acute, acuminate, or cuspidate, base attenuate, petioles 2-5 cm long, Inflorescences 2(-4)-flowered, peduncles 50-120 mm long, curved downward, pedicels 20-35 mm long, abruptly ascending; hypanthium hemispherical to obconical, 15-20 mm long, 15-20 mm wide; perianth blackish purple, 40-55 mm long, 15-20 mm wide, glabrous, the tube suberect to slightly curved, the lobes spreading, about as long as the tube; staminal column dark purple, glabrous or sparsely pubescent; anthers dark purple, glabrous or sparsely pubescent. Berries orange, subglobose, 20-30 mm long, 20-30 mm wide. [2n=28*.] Occurring in wet and low boggy forest, 915-1,460 m, on the summit of the Kohala Mountains, Hawai'i. — Plate 48.

5. Clermontia fauriei H. Lév.

[C. clermontioides (Gaud.) A. Heller var. epiphytica Hochr.; C. c. var .hirsutiflora Rock]

(end) Hāhā'aiakamanu

Terrestrial or epiphytic shrubs or trees 2-7 m tall. Leaves oblong to elliptic, blades 5-17 cm long, 1.5-7 cm wide, glabrous, upper surface dark green, glossy, lower surface pale green, margins callose-crenulate, apex acute, acuminate, or cuspidate base cuneate or obtuse, petioles 1.5-8 cm long. Inflorescences 1-2-flowered, glabrous or rarely pubescent, peduncules 8-30 mm long, pedicels 5-30 mm long; hypanthium hemispherical to obconical, 9-15 mm long, 10-11 mm wide, glabrous or rarely pubescent; calyx lobes deltate to shallowly triangular, 2-4 mm long, 2-4 mm wide, apex acute to acuminate; corolla greenish or purplish externally, cream within, 60-70 mm long, 5-8 mm wide, glabrous or pubescent, the tube arcuate, the lobes 1/4-1/5 as long as the tube, all curved downward in the same plane, the corolla thus appearing unilabiate; staminal column greenish white to cream, exserted, glabrous anthers cream-colored, glabrous or sparsely pubescent. Berries orange, subglobose, 25-35 mm long, 25-35 mm wide. Occurring in diverse mesic forest to wet forest, 365-1,400m, on Kaua'i and O'ahu. — Plate 49.

The name Clermontia clermontioides has long been applied to the sole representative of the genus on Kaua'i. However, examination of the type (Gaudichaud 147, P) revealed that this name is correctly applied to plants of Hawai'i previously called Clermontia coerulea. The species on Kaua'i must now be called Clermontia fauriei. Infraspecific taxa described from Maui (Clermontia clermontioides var. barbata) and Hawai'i (C. c. var. singuliflora) are here included in Clermontia peleana. Although long recognized as a common species on Kaua'i, C. fauriei has not previously been reported from O'ahu. Two collections are known from the extreme western end of that island, one from each mountain range. The earlier was collected at Kolao Gulch in the Ko'olau Mountains in 1950 (Degener 20397, B, BISH, BM, NY, W). Six years later, it was collected on the ridge east of central Makeleha Valley in the Wai'anae Mountains (Ozaki 1568, 1569, 1570, BISH).

page 429

page 430

6. Clermontia grandiflora Gaud.

[C. earina St. John; C. forbesii St. John; C. grandiflora var. brevifolia A. Gray, nom. illeg.; C. g. var forbesii (St. John) F. Wimmer; C. g. var. f. f. hirsutinervis (St. John) F. Wimmer; C. g. f. hamata F. Wimmer; C. g. subsp. munroi (St. John) Lammers; C. g. var. subpetiolata (St. John) F. Wimmer; C. g. var. vulgata F. Wimmer; C. g. var. v. f. nitida F. Wimmer; C. hirsutinervis St. John; C molokaiensis St. John; C. munroi St. John; C. reticulata St. John; C. r. f. pilifera St. John; C. spatulata St. John; C. subpetiolata St. John; C. wailauensis St. John; Delissea filigera Wawra; Lobelia grandiflora (Gaud.) Endl.]

(end)

Terrestrial or rarely epiphytic shrubs or trees 1-6m tall. Leaves elliptic or oblanceolate, blades 4-15 cm long 1.5-4.5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent, margins callose-crenulate, apex acute, acuminate, or cuspidate, base cuneate or attenuate, petioles 0.2-4 cm long. Inflorescences 2(-5)-flowered, pendulous, peduncles 20-110 mm long, pedicels 18-65 mm long; hypanthium turbinate or obovoid, 10-17 mm long, 10-20 mm wide; perianth greenish, purplish, maroon, or rose, curved to arcuate, distinctly tubular, 55-80 mm long, 7-20 mm wide, glabrous, the lobes (except the dorsal) erect or downcurved slightly, 1/5-1/3 as long as the tube; staminal column purple or magenta, glabrous; anthers purple, glabrous. Berries orange, obovoid to subglobose, 16-26 mm long, 10-20 mm wide. [2n=28*.] Occurring in wet forest and margins of bogs, 530-1,975 m, on Moloka'i, Lāna'i, and Maui. — Plate 49.

Two subspecies may be recognized: subsp. grandiflora [incl. Clermontia earina, C. grandiflora var. brevifolia, C. g. f. hamata, C. reticulata f. pilifera, Delissea filigera] with perianth curved, stout, 12-20 mm wide, 3-5 times longer than wide (Maui); and subsp. munroi [incl. C. forbesii. C. grandiflora var. vulgata, C. g. var. v. f. nitida, C. hirsutinervis, C. molokaiensis, C. reticulata, C. spatulata, C. subpetiolata, C. wailauensis] with perianth arcuate, slender, 7-10 mm wide, 5-9 times longer than wide (Moloka'i, Lāna'i, Maui). On West Maui, subsp. grandiflora is common at 760-1,675 m, while subsp. munroi is known from only a few specimens collected at 610-760m. On East Maui, no ecological or geographical separation is discernible from the available data.

7. Clermontia hawaiiensis (Hillebr.) Rock.

[C. macrocarpa Gaud. var. hawaiiensis Hillebr.; C. grandiflora Gaud. var. longifolia A. Gray; C. kohalae Rock var. hiloensis F. Wimmer; C. subteralbulus St. John]

(end) 'Ōhā kēpau, 'ōhā wai nui, 'Ōhā-hā wai nui

Terrestrial or epiphytic trees or shrubs 1.5-9 m tall. Leaves oblong or oblanceolate, blades 9-24 cm long, 2.5-6.5 cm wide, upper surface dark green, glossy, glabrous, lower surface whitish green, glabrous or pubescent along the veins, margins callose-crenulate, apex acuminate, acute, obtuse, or rounded, base attenuate, petioles 2-7 cm long. Inflorescences 2(-4)-flowered, peduncles 15-60 mm long, pedicels 15-40 mm long; hypanthium obconical, 17-22 mm long, 11-18 mm wide, prominently 10-ridged, the dorsal ridge forming a definite keel; perianth greenish white, sometimes tinged maroon apically, curved, distinctly tubular, 50-65 mm long, 9-18 mm wide, glabrous, the lobes (except the dorsal) erect or downcurved, 1/5-1/3 as long as the tube, prominently ridged externally; staminal column maroon or rarely greenish white, glabrous; anthers purple or rarely greenish white, glabrous or sparsely pubescent. Berries orange obovoid or ellipsoid, 10-ridged, 25-37 mm long, 15-30 mm wide. Occurring in mesic to wet forest, 550-1,760 m, Puna and Ka'ū districts, Hawai'i.

Reports from Moloka'i (Degener & Degener, 1958; Wimmer, 1953, 1968) are based upon a misidentified specimen (Degener 7826, NY) of Clermontia kakeana. Clermontia hawaiiensis occasionally hybridizes with C. parviflora (e.g. Lammers et al. 5814, BISH, OS). Although reported to hybridize with C. montis-loa (Carlquist, 1980), the voucher specimens (Carlquist 2025, BISH, C. DUKE, E, F, GH, L, MICH, RSA, UC, Z) also represent C. hawaiiensis x C. parviflora.

page 431

8. Clermontia kakeana Meyen

[C. glabra St. John; C. kakeana var. forbesii Degener & I. Degener; C. k. f. gracilis F. Wimmer; C. k. var. orientalis St. John; C. k. f. rosea (Hillebr.) Degener & I. Degener; C. macrocarpa Gaud.; C m. var. cymosa Hillebr.; C. m. var. rosea Hillebr; C. macrophylla Nutt.; C. montis-loa Rock f. molokaiensis F. Wimmer; Lobelia kakeana (Meyen) Endl.]

(end)

Terrestrial or epiphytic shrubs or trees 1-6 tall. Leaves elliptic, oblanceolate or narrowly obovate, blades 12-26 cm long, 3.5-9.8 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent, margins callose-crenulate, apex obtuse, acute, acuminate, or cuspidate, base cuneate to attenuate, petioles 2-7 cm long, glabrous or pubescent, Inflorescences 2(-7)-flowered, glabrous or sparsely pubescent, peduncles 15-55 mm long, pedicels 10-25 mm long; hypanthium hemispherical to obovoid, 8-14 mm long, 8-14 mm wide, glabrous or sparsely pubescent; perianth pale green externally, greenish white to white within, very rarely rose, 45-55 mm long, 8-14 mm wide, glabrous or sparsely pubescent, the tube suberect to curved, the lobes spreading, about as long as the tube; staminal column magenta or rarely white, glabrous; anthers dark purple, rarely white, glabrous. Berries subglobose to oblong, 18-34 mm long, 15-34 mm wide. [2n=28*.] Occurring in mesic forest, occasionally on margins of wet forest, 245-1,100 m, on O'ahu, Moloka'i, and Maui. — Plate 49.

A single plant believed to be a hybrid between this species and Clermontia micrantha was observed at Lanilili, West Maui, in 1984-1985 (Lammers & Sylva 5619, OS; Lammers 5847, BISH, OS, US). Clermontia grandiflora var. longifolia was placed in synonymy under C. kakeana by Hillebrand (1888), Rock (1919a), and Wimmer (1943). However, the type (U. S. Expl. Exped. s.n., US) is a specimen of Clermontia hawaiiensis.

9. Clermontia kohalae Rock

[C. convallis F. Wimmer; C. epilosa St. John; C. kohalae var. robusta Rock; C. leptoclada Rock var. holopsila F. Wimmer; C. l. var. urceolata Rock]

(end)

Terrestrial or epiphytic trees 2-6 m tall. Leaves narrowly elliptic or oblanceolate, blades 10-21 cm long, 2-5.5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or sparsely pubescent, margins callose-crenulate, apex obtuse, acute or acuminate, base attenuate, petioles 2-6 cm long. Inflorescences 2(-4)-flowered, peduncles 15-45 mm long, pedicels 15-35 mm long; hypanthium obconical or obovoid, 10-15 mm long, 10-15 mm wide; perianth blackish purple, 50-65 mm long, 9-15 mm wide, glabrous, the tube suberect or slightly curved, the lobes spreading, about as long as the tube; staminal column dark purple, glabrous; anthers dark purple, glabrous or sparsely pubescent. Berries orange subglobose, 20-32 mm long, 20-32 mm wide. Occurring in wet forest, 370-1,370 m, Kohala and Hāmākua districts, Hawai'i.

As suggested by Carlquist (1969a), Clermontia leptoclada Rock appears to represent a hybrid between C. kohalae and C. drepanomorpha. Clermontia convallis and C. leptoclada var. holopsila, unintentionally based on duplicates of the same specimen (Degener 2186, A, BISH, NY), represent typical Clermontia kohalae. The type of C. kohalae var. hiloensis ( Degenera 2190, A, B, BISH, NY, W) is a specimen of C. hawaiiensis.

10. Clermontia lindseyana Rock

[C. albimontis St. John; C. hawaiiensis (Hillebr.) Rock var. grandis Rock; C. linseyana var. livida Rock; C. viridis St. John]

(end, E*)

Terrestrial shrubs or trees 2.5-6 m tall. Leaves oblanceolate or elliptic, blades 13-24 cm long, 3.8-6.5 cm wide, upper surface dark green, glabrous or sparsely pubescent along the midrib, lower surface pale green or purplish, pubescent, margins callose-crenulate, apex acuminate, acute, or obtuse, base attenuate, petioles 2.5-7 cm long, pubescent. Inflorescences 2-flowered, pubescent, peduncles 25-40 mm long, pedicels 10-25 mm long; hypanthium hemispherical or obovoid, 12-20 mm long, 10-22 mm wide, pubescent; perianth green or rarely tinged purplish externally, white or cream within, 55-65 mm long, 9-18 mm wide, pubescent, the tube slightly curved, the lobes spreading, about as long as the tube; staminal column white, glabrous; anthers white, glabrous. Berries orange, subglobose, 25-40 mm long, 25-40 mm wide. Rare in mesic forest, 1,220-1,825 m, Hilo, Ka'ū, and South Kona districts, Hawai'i, and on the leeward slopes of Haleakalā, East Maui.

This species is similar to Clermontia kakeana, but differs in its larger, more pubescent flowers and more pubescent leaves. Its occurrence on Maui is documented by specimens from Kaupō (Rock 8688, BISH, GH, NSW) and Pu'upane (Forbes 1888.M, BISH, AS, P, UC, US). These specimens were included by St. John (1971a) in his Clermontia kakeana var. orientalis.

page 432

11. Clermontia micrantha (Hillebr.) Rock

[C. multiflora Hillebr. var. micrantha Hillebr.; C. m. var. m. f. montana Rock]

(end)

Terrestrial shrubs 0.4-2 m tall. Leaves elliptic to oblong, blades 4-10 cm long 1-2.5 cm wide, glabrous, margins callose-crenulate, apex acute or acuminate, base cuneate or attenuate, petioles 1.5-4 cm long. Inflorescences 2-3(-5)-flowered, peduncles 5-10 mm long, pedicels 7-14 mm long; hypanthium obovoid to turbinate, 5-8 mm long, 3-7 mm wide; perianth rose to dark pink, 18-28 mm long, 3-5 mm wide, glabrous, the tube suberect, the lobes recurved, about as long as the tube; staminal column rose to dark pink, glabrous; anthers purplish, glabrous. Berries orange, obovoid, 12-15 mm long, 9-10 mm wide Occurring in bogs and occasionally wet forest, 670-1,460 m, on Lāna'i and West Maui. — Plate 49.

Clermontia micrantha is rather common in bogs of West Maui (Pu'ukukui, Mount 'Eke, Lanilili). On Lāna'i, it is known only from 3 specimens (Munro 333, BISH; Munro 391, BISH, L, M; Munro 930, BISH, L) collected in Kumoa Gulch during 1927-1929. An individual transplanted from Lāna'i to Munro's garden on O'ahu died in 1935 (Munro 9, BISH). A single plant believed to be a hybrid between this species and Clermontia kakeana was observed at Lanilili in 1984-1985 (Lammers & Sylva 5619, OS; Lammers 5847, BISH, OS, US).

12. Clermontia montis-loa Rock

[C. m. f. globosa Rock]

(end)

Terrestrial or epiphytic trees or shrubs 2-5 m tall. Leaves oblanceolate or oblong, blades 8-19 cm long, 2.2-4.5 cm wide, upper surface dark green, glabrous, lower surface pale green or purplish, pubescent, margins callose-crenulate, apex acute, acuminate, or cuspidate, base attenuate, petioles 1.5-6 cm long. Inflorescences 2-3-flowered, peduncles 3-18 mm long, pedicels 3-18 mm long; hypanthium obconical or obovoid, 10-16 mm long, 8-11 mm wide; perianth greenish or purple externally, purple or blackish purple within, 38-50 mm long, 7-11 mm wide, glabrous, the tube suberect, the lobes spreading, as long as the tube or a little longer; staminal column purple, included within perianth, glabrous; anthers dark purple, glabrous. Berries orange, subglobose to ellipsoid, 25-30 mm long, 15-30 mm wide. [2n=28*.] Occurring in wet forest, 1,070-1,700 m, Hilo, Puna, and Ka'ū districts, Hawai'i.

Clermontia montis-loa hybridizes with C. parviflora (e.g., Lammers 5419, BISH, DUKE, MO, NY, OS, US). The type of C. montis-loa f. molokaiensis (Degener 7847, A NY) is a specimen of C. kakeana. The type of C. montis-loa var. tenuifolia (Skottsberg 707, BISH, GB, S) is a specimen of C. calophylla.

13. Clermontia multiflora Hillebr.

(end, Ex)

Shrubs 2-4 m tall. Leaves elliptic, blades 10-12.5 cm long, 2.5-3.2 cm wide, glabrous, margins callose-crenulate, apex acuminate, base cuneate, petioles 4-7.5 cm long. Inflorescences 7-10-flowered, peduncles 10-18 mm long, pedicels 10-18 mm long; hypanthium obconical, 7-8 mm long, 7-8 mm wide; perianth purplish, 24-32 mm long, 5-6 mm wide, glabrous, the tube suberect; staminal column glabrous; anthers purple, glabrous. Berries unknown. Known only from the syntype material collected in 1870 (Hillebrand s.n., BISH, K, MEL), on O'ahu and West Maui.

page 433

14. Clermontia oblongifolia Gaud.

[C. aspera F. Wimmer, pro parte; C. grandiflora Gaud. var. oblongifolia (Gaud.) A. Gray, pro parte; C. oblongifolia f. brevipes F. Wimmer; C. o. subsp.brevipes (F. Wimmer) Lammers; C. o. f. kaalae Degener; C. o. f. mauiensis (Rock) Degener; C. o. subsp. mauiensis (Rock) Lammers; C. o. var. mauiensis Rock; Lobelia oblongifolia (Gaud.) Endl.]

(end, 2subspp. E*)

Terrestrial shrubs or trees 2-7 m tall. Leaves oblong, elliptic, or lanceolate, blades 7-19 cm long, 2-5 cm wide, upper surface dark green, glossy, glabrous, lower surface whitish green, glabrous or pubescent, margins callose-crenulate, apex obtuse, acute, or acuminate, base cuneate to attentuate, petioles 1.8-11.5 cm long. Inflorescences 2(3)-flowered, peduncles spreading, 5-45 mm long, pedicels 10-45 mm long; hypanthium hemispherical to obconical, 10-13 mm long, 10-13 mm wide; perianth greenish white or purplish externally, white or cream within, arcuate, distinctly tubular, 50-65 mm long, 10-13 mm wide, glabrous, the lobes (except the dorsal) erect or spreading slightly, 1/4-1/3 as long as the tube; staminal column white or purple, glabrous; anthers white or purple, glabrous or sparsely pubescent. Berries orange, subglobose, 17-30 mm long. [2n=28*; the count of 2n=22* (Carlquist, 1956) is probably an error.] Occurring in mesic valleys to wet forest, 400-1,200 m, on O'ahu, Moloka'i, Lāna'i, and Maui.

Three subspecies may be distinguished. The first, subsp. brevipes, differs from the other 2 in its lanceolate leaf blades 7-11 cm long (versus oblong or elliptic, 8-19 cm long), petioles 1.8-3 cm long (versus 2.5-11.5 cm long), and peduncles 5-10 mm long (versus 10-45 mm long) (Moloka'i). Of the remaining 2, subsp. mauiensis is further characterized by its acuminate leaf apices, magenta or purple staminal column and anthers, and flower buds acuminate at the apex (Lāna'i, Maui). In contrast, subsp. oblongifolia [incl. Clermontia aspera, C. oblongifolia f. kaalae] has rounded, obtuse or acute leaf apices, greenish white or white staminal column and anthers, and flower buds rounded or obtuse at the apex (O'ahu). Clermontia aspera is based upon a mixed collection (Degener et al. s. n., W), comprising a detached inflorescence of Clermontia oblongifolia subsp. oblongifolia and leaves of Rollandia lanceolata subsp. calycina. The recently described Clermontia oblongifolia f. glabra (St. John, 1983b) is based upon a rather typical specimen (St. John et al. 23383, BISH) of C. pallida.

Although subsp. oblongifolia is still relatively common, the other 2 subspecies are much rarer. The Moloka'i race has been collected only once since 1938, in 1962 (Lamoureux 1990, HAW). Specimens of subsp. mauiensis were last collected on East Maui in 1927 (Degener 7947, GH, US), and on Lāna'i in 1913 (Forbes 44.L, BISH; Munro 55, BISH). Although not previously reported from West Maui, its occurrence there is documented by a few recent specimens, collected at 950-990 m along the trail to Pu'ukukui (e.g., Lammers & Hobdy 5690, OS).

15. Clermontia pallida Hillebr.
[C. oblongifolia Gaud. f. glabra St. John; C. pallida var. ramosissima Rock]
(end)

Terrestrial or rarely epiphytic shrubs or trees 2-5 m tall. Leaves oblong, elliptic, or oblanceolate, blades 7-15 cm long, 2-4.5 cm wide, glabrous, upper surface green, lower surface whitish green, margins callose-crenulate, apex acuminate to cuspidate, base cuneate, petioles yellowish white, translucent, 5-13.5 cm long. Inflorescences 2-3(-7)-flowered, peduncles 8-27(-50) mm long, pedicels 14-23(-30) mm long; hypanthium hemispherical, 7-10 mm long, 7-10 mm wide; perianth blackish purple to dark maroon externally, pale greenish within and at the tips of the lobes externally, 45-60 mm long, 7-10 mm wide, glabrous, the tube slightly curved, the lobes spreading, about a long as the tube; staminal column dark purple, glabrous or rarely minutely pubescent; anthers dark purple, glabrous. Berries unknown. Occurring in wet forest, 915-1,390 m, Moloka'i. — Plate 50.

page 434

page 435

16. Clermontia parviflora Gaud. ex A. Gray
[C. paradisia F. Wimmer; C. parviflora var. pleiantha Hillebr.; C. p. var. umbraticola Skottsb.; Cyanea blinii H. Lév.]
(end)

Terrestrial or epiphytic shrubs 1-3.5 m tall. Leaves elliptic or oblanceolate, blades 6-18 cm long, 1.5-5.5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or sparsely pubescent, margins callose-crenulate, apex acute, acuminate, or cuspidate, base cuneate or attenuate, petioles 1.5-5 cm long, glabrous or pubescent. Inflorescences (2)3-7-flowered, glabrous or pubescent, peduncles 10-15 mm long, pedicels 8-10 mm long; hypanthium turbinate or obovoid, 5-8 mm long, 3-5 mm wide, glabrous or sparsely pubescent; perianth green, purple, or white externally white or pale purple within, 15-28 mm long, 3-5 mm wide, glabrous or rarely minutely pubescent, the tube suberect, the lobes recurved, as long as the tube or a little longer; staminal column white or pale purple, exserted; anthers white or pale purple, glabrous. Berries yellow or orange oblong or subglobose, 12-25 mm long, 10-20 mm wide. [2n=28*.] Occurring in wet forest, 120-1,460 m, Kohala Mountains and windward slopes of Mauna Kea and Mauna Loa, Hawaii. — Plate 50.

This shrub is quite promiscuous, hybridizing with Clermontia hawaiiensis in Puna (e.g. Lammers et al. 5814, BISH, OS); C. montis-loa on windward Mauna Loa and Mauna Kea (e.g., Lammers 5419, BISH, DUKE, MO, NY, OS, US); and C. kohalae in the Kohala Mountains (e.g. Degener 21881, B, BISH, MASS, W). The last-cited specimens were among those used by Wimmer (1953) in describing C. paradisia. However, the specimen designated as lectotype (Degener 21880, B, BISH, MASS, NY, W) by Degener and Degener (1956) represents typical Clermontia parviflora, and C. paradisia is thus treated as a synonym of that species. Clermontia parviflora var. grandis and C. p. var. intermedia are here treated as synonyms of C. calophylla, while C. p. var. calycina is treated as a synonym of C. waimeae.

17. Clermontia peleana Rock
[C. clemontioides (Gaud.) A. Heller var. barbata (Rock) St. John; C. c. var. mauiensis Hochr., nom. illeg.; C. c. var. singuliflora (Rock) Hochr.; C. gaudichaudii Hillebr. var. barbata Rock; C. g. var. singuliflora Rock; C. singuliflora (Rock) Rock]
(end, E*)

Epiphytic shrubs or trees 1.5-6 m tall. Leaves oblong to elliptic, blades 8-20 cm long, 3-5 cm wide, glabrous, margins callose-crenulate, apex acute to acuminate, sometimes nearly cuspidate, base cuneate, obtuse, or almost rounded, petioles 3-6 cm long. Inflorescences 1-2-flowered, peduncles 8-17 mm long, pedicels 30-45 mm long; hypanthium obconical, 10-15 mm long, 10-15 mm wide; calyx lobes triangular, 1-2 mm long, 0.5-1 mm wide; corolla blackish purple or greenish white, 50-70 mm long, 8-13 mm wide, glabrous, the tube arcuate, the lobes 1/4-1/5 as long as the tube, all downcurved in the same plane, the corolla thus appearing unilabiate; staminal column exserted, glabrous; anthers glabrous or sparsely pubescent at apex. Berries orange, subglobose, 25-30 mm long. Extremely rare, epiphytic on 'ōhi'a, koa, 'ōlapa, and tree ferns in wet forest, 530-1,150 m, on East Maui and windward slopes of Mauna Kea and Mauna Loa, Hawai'i. — Plate 50.

18. Clermontia persicifolia Gaud.
[C. epiphytica St. John; C. grandiflora Gaud. var. oblongifolia (Gaud.) A. Gray, pro parte; Lobelia persicifolia (Gaud.) Endl., non Lam. nec Cav. nec Kunth]
(end)

Terrestrial or epiphytic shrubs or trees 2-6 m tall. Leaves elliptic to oblanceolate, blades 7-16 cm long, 1.5-4 cm wide, glabrous, upper surface dark green, glossy, lower surface pale green, margins callose-crenulate, sometimes revolute, apex acute, obtuse, or rounded, rarely acuminate, base cuneate or attenuate, petioles 2-7 cm long. Inflorescences 2-3(-6)-flowered, peduncles 6-30 mm long, pedicels 12-30 mm long; hypanthium hemispherical, obconical, or oblong, 7-12 mm long, 6-8 mm wide; perianth greenish white to white, sometimes with a faint purplish tinge externally, 48-60 mm long, 5-8 mm wide, glabrous, the tube suberect to gently curved, the lobes spreading, about as long as the tube; staminal column white, glabrous; anthers white, glabrous. Berries yellowish orange, subglobose, 17-30 mm long. Occurring in wet forest, 305-850 m. O'ahu, much more common in the Ko'olau Mountains than the Wai'anae Mountains.

Reports from Lāna'i (Rock, 1919a; Wimmer, 1943) are based upon a misidentified specimen (Hitchcock 14643, US) of Clermontia grandiflora subsp. munroi.

page 436

19. Clermontia pyrularia Hillebr.
[Delissea obtusa A. Gray var. mollis A. Gray]
(end, E*)

Terrestrial trees 3-4 m tall. Leaves narrowly elliptic or rarely oblanceolate, blades 5-28 cm long, 2.5-5 cm wide, upper surface green, dull, glabrous or sparsely pubescent, lower surface pale green, pubescent, margins callose-serrulate, apex acute to acuminate, mucronulate, base attenuate, narrowing imperceptibly into a winged petiole 1.5-3.5 cm long, Inflorescences 2-3(-5)-flowered, sparsely pubescent, peduncles 28-60 mm long, pedicels 8-20 mm long; hypanthium obconical to obovoid, 10-18 mm long, 8-12 mm wide, pubescent; calyx lobes triangular, 3-5 mm long, 1-2 mm wide, pubescent, apex acute or acuminate; corolla white or greenish white, 40-45 mm long, 5-8 mm wide, pubescent, the tube curved, the lobes spreading, about as long as the tube or a little shorter; staminal column glabrous; anthers glabrous. Berries orange, obovoid to obpyriform, 18-28 mm long, 15-24 mm wide. Rare in mesic to wet forest, 1,585-2,130 m, on windward Mauna Kea and leeward Mauna Loa, Hawai'i.

Delissea obtusa var. mollis was based upon a sterile specimen (U.S. Expl. Exped. s.n., US) of Clermontia pyrularia collected on Mauna Kea, Hawai'i. Hillebrand (1888) and Rock (1919a), neither of whom saw the type, treated the name as a synonym of Cyanea obtusa, a species otherwise restricted to East Maui.

20. Clermontia samuelii C. Forbes
[C. gracilis St. John; C. hanaensis St. John; C. kipahuluensis St. John; C. rosacea St. John; C. samuelii subsp. hanaensis (St. John) Lammers]
(end)

Terrestrial shrubs 1.2-5 m tall. Leaves oblanceolate to elliptic or broadly elliptic, blades 5-10 cm long, 1.8-4.5 cm wide, upper surface dark green, often tinged purplish, glabrous or sparsely pubescent, lower surface pale green, sparsely to densely pubescent, margins callose-crenulate, apex acute, acuminate, or cuspidate, base attenuate, cuneate, or obtuse, petioles 0.5-2.5 cm long. Inflorescences 2(-5)-flowered, peduncules 4-18 mm long, pedicels 12-28 mm long; hypanthium obconical, 8-14 mm long, 5-10 mm wide; perianth rose or greenish white to white, curved, distinctly tubular, 36-55 mm long, 5-10 mm wide, glabrous, the lobes erect, 1/5-1/3 as long as the tube; staminal column glabrous; anthers glabrous. Berries unknown. Occurring in wet forest and margins of bogs, 610-2,100 m, Hāna District, East Maui.

Two subspecies may be recognized: subsp hanaensis [incl. C. kipahuluensis] with perianth greenish white to white, 40-55 mm long, and blades broadly elliptic, 1.5-2.5 times longer than wide, lower surface sparsely pubescent (Kūhiwa Gulch to Pu'uiki, 610-915 m); and subsp. samuelii [incl. Clermontia gracilis, C. rosacea] with perianth rose, 36-48 mm long, and blades oblanceolate to elliptic, 3-4 times longer than wide, lower surface densely pubescent (Kīpahulu to Wai'ānapanapa, 1675-2100 m).

21. Clermontia tuberculata C. Forbes
[C. t. var. subtuberculata St. John]
(end)

Terrestrial shrubs or trees 4-5 tall. Leaves oblanceolate, oblong, or obovate, blades 9-20 cm long, 2.5-5.5 cm wide, upper surface glabrous, lower surface glabrous or minutely muricate on the veins, margins callose-crenulate, apex obtuse, base cuneate or attenuate, petioles 1-6 cm long. Inflorescences 1-2-flowered, muricate, peduncules 5-10 mm long, pedicels 30-55 mm long; hypanthium obconical or obovoid, 14-19 mm long and wide, tuberculate; calyx lobes triangular, deltate, or shallowly triangular, 3-9 mm long, apex obtuse or acute, tuberculate; corolla rose, 50-70 mm long, 12-18 mm wide, tuberculate, the tube suberect, the lobes thick, spreading, as long to twice as long as the tube; staminal column magenta, glabrous; anthers dark purple, glabrous or sparsely pubescent. Berries subglobose, muricate. Occurring in wet forest, primarily along streambeds, 1,650-1,825 m, East Maui. — Plate 50.

This species is closely related to Clermontia arborescens, differing primarily in the muricate inflorescences, rose corolla, and more deeply cleft lip of the corolla. Although many species of Cyanea bear epidermal murications on flowers, leaves, and stems (Carlquist, 1962a), this is the only such species of Clermontia. Perhaps not coincidentally, C. tuberculata is entirely sympatric with Cyanea aculeatiflora. The latter is closely related to Cyanea macrostegia, differing primarily in characters similar to those separating Clermontia arborescens and C. tuberculata; muricate inflorescences and more deeply cleft corolla.

page 437

22. Clermontia waimeae Rock
[C. bicolorata St. John; C. kahuaensis St. John; C. parviflora Gaud. ex A. Gray var. calycina Rock; C. waimeae var. longisepala Rock; C. w. var. l. f. lanceolata F. Wimmer; C. w. var. obovata Rock]
(end)

Terrestrial or epiphytic shrubs or trees 1-7 m tall. Leaves oblong, elliptic, or oblanceolate, blades 5-15 cm long, 2-5.5 cm wide, upper surface glabrous, lower surface glabrous or pubescent, margins callose-crenulate, apex obtuse, acute, acuminate, or rarely cuspidate, base cuneate, petioles 1-5 cm long. Inflorescences 2-3(-5)-flowered, peduncles 8-30 mm long, pedicels 5-18 mm long; hypanthium hemispherical, obconical, or oblong, 8-12 mm long, 6-10 mm wide; calyx lobes triangular to narrowly triangular, 3-25 mm long, 0.5-3 mm wide, apex acute to acuminate; corolla blackish purple at least on the lobes, greenish on the tube, 30-40 mm long, 4-6 mm wide, glabrous or sparsely pubescent, the tube suberect, the lobes spreading, about as long as the tube; staminal column purple, glabrous; anthers purple, glabrous or sparsely pubescent. Berries orange, subglobose, 20-25 mm long. Occurring in bogs and wet forest, 1,070-1,520 m, Kohala Mountains, Hawai'i. — Plate 51.

3. CYANEA Gaud. Hāhā

[Delissea Gaud. sect. Cyanea (Gaud.) Baill.; Kittelia Rchb., nom. illeg.; Macrochilus K. Presl]

Branched or unbranched shrubs or trees 0.3-6 m tall, sometimes palm-like and 1-14 m tall, terrestrial or rarely epiphytic, with white, yellow, or tan latex; stems unarmed, muricate, or aculeate. Leaves callose-toothed or pinnately lobed, cleft, parted, or divided, the lobes or segments then callose-toothed, lobed or divided. Flowers 5-25(-40) in axillary racemes, the rachis sometimes not expanded, the raceme thus appearing subumbellate; hypanthium obconical or obovoid, rarely ovoid, oblong, or depressed-obovoid; calyx lobes dentiform, triangular, or oblong, rarely linear, distinct or rarely connate, much shorter to somewhat longer than hypanthium, rarely up to as long as the corolla; corolla zygomorphic, glabrous or pubescent, rarely muricate, the tube suberect to strongly arcuate, dorsally cleft to about the middle, the lobes spreading, rarely erect or all 5 down-curved in the same plane; stamens connate; anthers with apical tufts of white hairs on the purple berry. Seeds brown to black, smooth, shiny, 0.2-0.5(-2) mm long. [Chamisso, 1833a; Degener & Degener, 1982; Forbes, 1912, 1916; Forbes & Munro, 1920; Gaudichaud-Beaupré, 1829a; Gray, 1861b; Heller, 1897; Hosaka & Degener, 1938; Léveillé, 1911b, 1913b; Meyen, 1843; Rock, 1915, 1917b, 1918a, 1957, 1962a; St. John, 1939b, 1969b, 1971a, 1975b, 1976a, 1978c, 1978d, 1980d, 1980e, 1981d, 1985a, 1987b, 1987c, 1987d; St. John & Takeuchi, 1987; Skottsberg, 1926, 1927b, 1944a; Swanholm, St. John & Scheuer, 1959, 1960; Wawra, 1873d]

Cyanea, with 52 species, is the largest genus of Hawaiian Campanulaceae, and among the largest genera of Hawaiian plants. Name derived from the Greek cyaneos, blue, in reference to the supposedly blue flowers of the type species, Cyanea grimesiana.

page 438

page 439

The genus has been divided into 3 (Hilleband, 1888; Wimmer, 1943; St. John, 1969b) or 5 (Rock, 1919a; Stone, 1967b) sections. Neither arrangement satisfactorily reflects relationships within this complex genus. In several species (e.g., Cyanea platyphylla, C. tritomantha) dramatic morphological changes are associated with development (Carlquist, 1962a, 1969a, 1974, 1980). Generally, juveniles of such species are more densely armed and bear more deeply lobed or dissected leaves. As development progresses, stems and leaves become less armed and leaf margins less lobed or dissected. Some species may flower precociously, while others are not known to do so. They key to species emphasized floral characters in order to minimize difficulties caused by this developmental variation. Generally, juvenile features are included in the descriptions only if the species is known to flower precociously.

As this book was going to press, several papers affecting the taxonomy of Cyanea appeared in a non-refereed journal. In the first, St. John and Takeuchi (1987) questioned the "validity" of the characters used by Hillebrand (1888), Rock (1919a), and others to distinguish Cyanea from Delissea. Acting on their conclusions, St. John (1987b) formally merged the 2 genera, publishing 103 new names and new combinations in Delissea, the genus with priority (St. John, 1987b). All of these are accounted for in the present treatment. This regrettable proliferation of names resulted from the wholesale transfer of every basionym ever treated as a Cyanea into Delissea. Apparently, types were not studies, nor was any attempt made to critically evaluate the taxa transferred. This merger is not accepted here on the grounds that it fails to provide a more natural classification of the plants involved, and is based upon an inadequate understanding of generic limits in the subfamily as a whole. The Lobelioideae, like the Asteraceae (cf. Lane & Turner, 1985), is a natural group in which nearly all of the genera are very poorly defined (Gleason, 1925; Moeliono & Tuyn, 1960; McVaugh, 1965). If one were to insist on iron-clad generic distinctions among Lobelioideae, the entire subfamily might be reduced to a single, conceptually useless genus. Furthermore, Cyanea and Delissea are part of a monophyletic group that also includes Clermontia and Rollandia (Rock, 1919a; Lammers & Stein, in prep.) Clermontia, Delissea, and Rollandia are unquestionably natural groups. Heterogenous as to each other. However, merging it with Delissea does nothing to relieve this apparent unnaturalness. The net result is the creation of a larger and even more unnatural genus. To achieve a more natural classification, one could merge all 4 genera into a single large genus or divide Cyanea into several segregates. Until such time as careful and detailed study of diverse data permits a more natural realignment, it is deemed best to maintain the traditional circumscriptions of these genera.

In a third paper (St. John, 1987c), the expanded Delissea was further enlarged by the description of 50 new species and 4 new infraspecific taxa. Of these, 45 were based on specimens collected at just 6 localities in Hanalei District, Kaua'i. That so small an area should yield so many novelties staggers the imagination, and suggests that most might be be accommodated within previously known species. Unfortunately, time did not permit procurement and examination of types. Further, the so-called "diagnoses" are of little use in distinguishing these novelties from each other or from previously described species. The characters used are of minimal utility in the systematics of lobelioids, while critical characters (e.g., width and curvature of the corolla tube, relative length and posture of corolla lobes, size and shape of the hypanthium) are ignored. This woeful inadequacy is compounded by the lack of accompanying illustrations. In this regard, St. John's (1980e) comments regarding the Hawaiian novelties of Hector Léveillé seem particularly appropriate. For these reasons, it was regrettably impossible to account for all these novelties in the previously described species. The majority are probably referrable to Cyanea coriacea, C. fissa, and C. sylvestris, the common species of Hanalei.

page 440

KEY TO THE SPECIES OF CYANEA

page 441

page 442

page 443

1. Cyanea aculeatiflora Rock

[C. solenocalyx Hillebr. var schizocalyx Hillebr.; Delissea aculeatiflora (Rock) St. John]

(end)

Palm-like trees 2-7 m tall. Leaves oblanceolate, oblong, or elliptic, blades 40-100 cm long, 10-21 cm wide, upper surface glabrous or sparsely muricate on the veins, lower surface pubescent, muricate on the veins, margins callose-denticulate, apex rounded or obtuse, base obtuse or cuneate, petioles 10-18 cm long, muricate. Inflorescences 5-10-flowered, clustered among the leaves, muricate, peduncles 40-100 mm long, pedicels 10-15 mm long; hypanthium obconical, 14-22 mm long, 8-15 mm wide, muricate; calyx lobes oblong, 10-25 mm long, 3-8 mm wide, apex obtuse or truncate, muricate; corolla blackish purple externally, lobes greenish toward apex white or pale lilac within, 60-85 mm long, 7-11 mm wide, muricate externally, glabrous within, the tube curved or suberect, the lobes spreading, as long as the tube; staminal column magenta, glabrous; anthers dark purple, glabrous, the lower 2 with apical tufts of white hairs. Berries yellow to yellowish orange, obovoid or ellipsoid, 20-25 mm long, 17-20 mm wide, crowned by the persistent calyx lobes, muricate. Occuring in wet forest, 945-1,450 m, on the windward slopes of Haleakalā, East Maui. — Plate 51.

Cyanea aculeatiflora is closely related to C. macrostegia, differing primarily in its muricate inflorescences and more deeply cleft corolla. The type of Cyanea macrostegia Hillebr. var viscosa Rock (Rock 8791, A, BISH, CAS, GH, NY, US) may be a hybrid between these 2 species.

page 444

2. Cyanea acuminata (Gaud.) Hillebr.

[Delissea acuminata Gaud.; Cyanea acuminata var. calycina Hosaka; C. a. f. latifolia (Wawra) F. Wimmer; C. occultans St. John; Delissea acuminata var calycina (Hosaka) St. John; D. a. f. latifolia (Wawra) St. John; D. a. var. latifola Wawra; D. occultans (St. John) Degener & I.Degener; Lobelia acuminata (Gaud.) Hook. & Arnott, non Sw.]

(end)

Shrubs 0.3-2 m tall; stems subherbaceous, woody only at base, unbranched or rarely sparingly branched from base. Leaves oblanceolate to narrowly obovate or elliptic, blades 11-32 cm long, 3-9 cm wide, upper surface green, glabrous or sparsely pubescent, lower surface whitish green, pubescent at least on the midrib and veins, margins minutely callose-denticulate, apex narrowly acuminate, base attenuate to cuneate, petioles 2-10 cm long. Inflorescences 6-20-flowered, pubescent, peduncles 15-60 mm long, pedicels 6-10 mm long; hypanthium obovoid, 4-5 mm long; calyx lobes narrowly triangular, 2-5 mm long, apex acuminate; corolla white, sometimes tinged purplish, 30-35 mm long, 3-4 mm wide, glabrous or sparsely pubescent, the tube suberect to slightly curved, the lobes spreading, 1/4-1/3 as long as the tube; staminal column glabrous or sparesely pubescent; anthers pubescent or rarely glabrous, the lower 2 with apical tufts of white hairs. Berries yellow to yellowish orange, globose, 4-6 mm long. Occurring in mesic valleys, 300-730 m, Ko'olau Mountains, O'ahu. — Plate 51.

3. Cyanea angustifolia (Cham.) Hillebr.

[Lobelia angustifolia Cham.; Cyanea angustifolia var. racemosa Hillebr.; Delissea acuminata Gaud. var. angustifolia (Cham.) A. Gray; D. angustifolia (Cham) K. Presl; D. a. var racemosa (Hillebr.) St. John; D. honoluluensis Wawra]

(end)

Branched shrubs 1.5-5 m tall. Leaves lanceolate, narrowly elliptic, or narrowly oblong, blades 9-32 cm long, 1-5-5.5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent along the midrib, margins callose-crenulate to callose-crenate, apex acuminate, base cuneate to attenuate, petioles 2.5-9 cm long. Inflorescences 6-25-flowered, spreading or pendent, peduncles 15-240 mm long, rachis elongate, up to 140 mm long, rarely condensed or almost obsolete, pedicels 8-30 mm long; hypanthium purple, obovoid, 3-6 mm long; calyx lobes dentiform, 0.5-1 mm long, pubescent at least at apex and margins, apex acute to acuminate; corolla white to greenish white, sometimes tinged pale purplish, 20-30 mm long, 2-3 mm wide, glabrous, the tube suberect to gently curved, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 5-10 mm long. [2n=28*; the count of 2n=24* (Carlquist, 1956) is probably an error.] Occurring in mesic forest and valleys to wet forest, 150-760 m, common in the Ko'olau Mountains, O'ahu, but rather scarce in the Wai'anae Mountains, and on eastern Moloka'i, Lāna'i, and northern West Maui. — Plate 51.

The infraspecific taxa described from Maui and Lāna'i (Cyanea angustifolia f. elliptica, C. a. var hillebrandii, C. a. var isabella, and C. a. var lanaiensis) are here treated as Cyanea elliptica, while C. angustifolia var. tomentella from the Wai'anae Mountains, O'ahu, is included in Cyanea membranacea.

4. Cyanea arborea Hillebr.

[Delissea arborea H. Mann, non K. Presl; Cyanea longifolia A. Heller, nom. illeg.]

(end, Ex)

Palm-like trees 4-8 m tall, with yellow latex. Leaves oblanceolate, 65-90 cm long, 6-12 cm wide, upper surface glabrous, lower surface sparsely pubescent, margins minutely callose-denticulate, undulate toward base, apex acute, base cuneate, sessile. Inflorescences 15-25-flowered, pendent, hanging below the leaves, peduncles 150-320 mm long, pedicels 4-8 mm long; hypanthium obovoid, 8-12 mm long; calyx lobes dentiform, 0.5-1 mm long; corolla cream tinged with pale lilac, 40-50 mm long, 3-5 mm wide, suberect to slighly curved, all 5 lobes curved downward in the same plane, the corolla thus appearing unilabiate, glabrous; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries globose, 10-12 mm in diameter. Formerly occurring in mesic forest, 1,520-1,650 m, leeward

page 445

slopes of Haleakalā, East Maui. Last collected in 1928 (Munro 804, A, BISH, DS, P).

The poorly known Cyanea arborea var. pycnocarpa from Kohala, Hawai'i, is here accepted as a distinct species, Cyanea pycnocarpa.

5. Cyanea asarifolia St. John

[Delissea asarifolia (St. John)St. John]

(end, E*)

Sparingly branched shrubs 0.3-1 m tall. Leaves cordiform, blades 8.5-10.5 cm long, 7-8 cm wide, glabrous, margins callose-crenulate, apex acute, base cordate, petioles 12-15 cm long. Inflorescences 30-40-flowered, glabrous, peduncles 25-30 mm long, pedicels 7-10 mm long; hypanthium obconical, 7-9 mm long; calyx lobes dentiform, 0.5-1 mm long; corolla white with purple longitudinal stripes, 20-22 mm long, 3-3.5 mm wide, gently curved, the lobes spreading; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Berries dark purple, globose ca. 1. cm long. Known only from the type (Hobdy 181, A, BISH), collected in 1970 on a sheer rock cliff, ca. 330 m, above the bed of Anahola Stream, northeastern Kaua'i.

6. Cyanea asplenifolia (H. Mann)Hillebr.

[Delissea asplenifolia H. Mann]

(end, Ex)

Shrubs 1.3-2 m tall; stems unbranched or sparingly, branched from base, sparsely muricate. Leaves pinnately divided, cut 3/4-7/8 the distance to the midrib, blades 22-42 cm long, 8-25 cm wide (across the segments), upper surface glabrous, lower surface sparsely pubescent and muricate on midrib and veins, the segments 10-15 per side, lanceolate to elliptic, 10-25 mm wide, margins of the segments irregularly lobed, the lobes obtuse, petioles 4.5-10 cm long, muricate. Inflorescences 8-15-flowered, peduncles 20-50 mm long, pedicels 8-12 mm long; hypanthium obconical, 4-6 mm long, 3-4 mm wide; calyx lobes narrowly oblong, 2-5 mm long, 1-2 mm wide, apex acute to acuminate; corolla 30-38 mm long, 3-5 mm wide, sparsely pubescent, the tube suberect to gently curved, the lobes spreading, 1/3-1/2 as long as the tube, sparsely muricate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Formerly occurring in mesic valleys, northern West Maui. Last collected in 1920 (Forbes 2540.M, L, WELT).

7. Cyanea comata Hillebr.

[Delissea comata (Hillebr.)St. John]

(end, Ex)

Sparingly branched shrubs 1.6-2.6 m tall. Leaves oblanceolate to narrowly obovate, blades 15-30 cm long, 5.5-9 cm wide, upper surface glabrous, lower surface sparsely pubescent, margins callose-denticulate, apex acute to obtuse and mucronate, base cuneate, petioles 2-4 cm long. Inflorescences 6-12-flowered, pendent, peduncles 150-300 mm long, pedicels 16-20 mm long, upcurved; hypanthium depressed-obovoid, 4-6 mm long, 6-8 mm wide, glabrous; calyx lobes triangular to deltate, 1-2.5 mm long; corolla pale lilac, 48-55 mm long, 3-5 mm wide, strongly arcuate, glabrous, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries depressed-obovoid, 6-8 mm long, 12-14 mm wide. Known only from a single set of 19th century specimens (Hillbrand 89, K; Hillebrand 100, B, K; Hillebrand s.n., BISH, BM, C, E, GH, MEL, NSW, S) collected at 915-1220 m on the leeward slopes of Haleakalā, East Maui.

8. Cyanea copelandii Rock

[C. c. subsp. haleakalensis (St. John) Lammers; C. haleakalaensis St. John; Delissea copelandii (Rock) St. John; D. haleakalaensis (St. John) St. John]

(end, [1 subsp. E*])

Liana-like shrubs 0.3-2 m tall, with tan latex; stems subherbaceous, reclining on moss-covered tree trunks and rooting at the nodes, or epiphytic, unbranched or sparingly branched from base. Leaves oblanceolate to narrowly or broadly elliptic, blades 10-27 cm long, 3.5-8.5 cm wide, upper surface glabrous, lower surface pubescent, margins callose-denticulate, apex narrowly acuminate to subcuspidate, base

page 446

cuneate to rounded, petioles 2.5-10 cm long. Inflorescences 5-12-flowered, pubescent, peduncles 20-45 mm long, pedicels 4-16 mm long; hypanthium obovoid, 6-10 mm long, 5-6 mm wide, pubescent; calyx lobes deltate to narrowly triangular, 2-4 mm long; corolla yellowish but appearing pale rose due to the dark red pubescence, 37-42 mm long, 4-5 mm wide, the tube gently curved, the lobes spreading, 1/4 as long as the tube or less; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries dark orange, ovoid, 7-15 mm long. Occurring in wet forest, 660-1,600 m, East Maui and Hawai'i. — Plate 52.

Two subspecies may be distinguised: subsp. copelandii with leaves narrowly elliptic to oblanceolate, 20-27 cm long (windward slopes of Mauna Loa, 660-880 m, Hawai'i); and subsp. haleakalaensis with leaves broadly elliptic, 10-19 cm long (windward slopes of Haleakalā, 1,300-1,600 m, East Maui). The former subspecies has been collected only one since 1918, in 1957 (Collector unknown s.n., BISH), and is apparently endangered.

9. Cyanea coriacea (A. Gray) Hillebr.

[Delissea coriacea A. Gray; D. c. var lumahaiensis St. John]

(end)

Unbranched or sparingly branched shrubs 2.5-3 m tall. Leaves oblong to oblanceolate, blades 20-38 cm long, 6-12 cm wide, glabrous, upper surface green, lower surface whitish green, margins minutely callose-crenulate, apex rounded, base cuneate, petioles 5-16 cm long. Inflorescences 12-20-flowered, the rachis condensed or obsolete, peduncles 25-80 mm long, pedicels 12-18 mm long; hypanthium obovoid, 4-5 mm long, 3-4 mm wide; calyx lobes dentiform, 0.5-1 mm long, pubescent on margins, apex acute to acuminate; corolla white tinged purplish, 20-25 mm long, 2-3 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 6-10 mm long. Occurring in mesic valleys, 150-220 m, northern Kaua'i. — Plate 52.

The types of Cyanea coriacea var. degeneriana (Degener 21737,B, W) and C. c. f. gratiosa (Degener 21736a, B, W) are specimens of Cyanea fauriei. The type of C. coriacea var. serratifolia (Webster 1636, BISH, MICH, US) is a specimen of C. membranacea

10. Cyanea degeneriana F. Wimmer

[C. bondiana (Rock) Rock; C. densiflora (Rock) Rock; C. floribunda F. Wimmer; C. pilosa A. Gray var. bondiana Rock; C. p. var. densiflora Rock; C. p. var.glabrifolia Rock; Delissea bondiana (Rock) St. John; D. degeneriana (F. Wimmer) St. John; D. densiflora (Rock) St. John; D. floribunda (F. Wimmer) St. John]

(end)

Shrubs 0.3-1.2 m tall, with tan latex; stems subherbaceous, woody only at base, unbranched or sparingly branched from base, pubescent. Leaves elliptic, blades 15-27 cm long, 4-8.5 cm wide, upper surface green, glabrous or sparsely pubescent, lower surface whitish green, pubescent at least on the midrib and veins, margins minutely callose-denticulate, apex narrowly acuminate, base cuneate, petioles 1-4 cm long. Inflorescences 6-16-flowered, pubescent, peduncles 5-20 mm long, pedicels 3-6 mm long; hypanthium obovoid, 4-5 mm long; calyx lobes narrowly triangular, 3-5 mm long, apex acuminate; corolla white, sometimes tinged with purple, 20-25 mm long, 3-4 mm wide, sparsely pubescent or subglabrous, the tube suberect to gently curved, the lobes spreading, 1/3-1/2 as long as the tube; staminal column glabrous; anthers pubescent, the lower 2 with apical tufts of white hairs. Berries orange, ellipsoid, 8-10 mm long. [2n=28*.] Occurring in wet forest, 460-1,520 m, Kohala Mountains, windward Mauna Kea, and windward Mauna Loa, Hawai'i. — Plate 52.

Rock (1913b) described 4 varieties of Cyanea pilosa (C. p. var. bondiana, C. p. var. densiflora, C. p. var. glabrifolia, and C. p. var. megacarpa), which were later (Rock, 1962a) recognized as distinct species. In the present treatment, the last is placed in synonymy under Cyanea pilosa subsp. pilosa, while the remainder are treated as comprising a single species. The type of Cyanea degeneriana (Park s.n., BISH) also is referrable to the latter.

page 447

page 448

Because this name is the oldest available, it must be used for the species as here circumscribed.

11. Cyanea dunbariae Rock

[Delissea dunbariae (Rock) St. John]

(end, Ex)

Branched shrubs 1.5-2 m tall; stems smooth. Leaves ovate to broadly elliptic, blades 10-22 cm long, 6-14 cm wide (across the lobes), margins irregularly lobed or cleft, apex acute to acuminate, base obtuse to truncate, petioles 4-12 cm long. Inflorescences 6-8-flowered, peduncles 30-70 mm long, pedicels 12-15 mm long; hypanthium ovoid, 4-6 mm long, 3-4 mm wide; calyx lobes triangular, 2-3 mm long, ca. 1 mm wide, apex acuminate; corolla white tinged or striped with pale lilac, 30-38 mm long, 3-4 mm wide, glabrous, the tube slightly curved, the lobes spreading, 3/4 as long to as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Known only from the type [Rock 13119 (coll. Dunbar), BISH, L] and 2 sterile specimens (Rock 13115, 14038, BISH) collected in 1918 at Waihānau, Moloka'i.

This poorly known species is similar to Cyanea profuga, another poorly known species endemic to Moloka'i. The 2 entities may represent geographic races or juvenile and adult phases of one species. Until more material is available for study, the status quo is maintained.

12. Cyanea elliptica (Rock) Lammers

[C. angustifolia (Cham.) Hillebr. f. elliptica Rock; C. a. var. elliptica (Rock) F. Wimmer; C. a. var Hillebrandii Rock; C. a. var isabella F. Wimmer; C. a. var lanaiensis Rock; C. a. var l. f. elliptica (Rock) St. John; Delissea angustifolia (Cham.) K. Presl var. hillebrandii (Rock) St. John; D. a. var. isabella (F. Wimmer) St. John; D. a. var. lanaiensis (Rock) St. John; D. a. var. l. f. elliptica (Rock) St. John; D. undulata Gaud. var. serrulata Wawra]

(end)

Branched shrubs 1-4 m tall. Leaves elliptic to oblong, blades 8-18 cm long, 2.8-7 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent along the midrib, margins callose-crenulate to callose-serrulate, apex obtuse to acute, base rounded to obtuse, petioles 2.5-6.5 cm long. Inflorescense 6-14-flowered, the rachis condensed or obsolete, peduncles 10-50 mm long, pedicels 5-20 mm long; hypanthium obovoid, 3-4 mm long, 2-3 mm wide; calyx lobes dentiform, 0.5-1 mm long, pubescent at the margins, apex acute to acuminate; corolla greenish white or white tinged purplish, 15-22 mm long, 1.5-3 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 6-9 mm in diameter. Occuring in mesic valleys and forest, 475-915 m, Lāna'i and Maui.

Cyanea elliptica was formed by merging 4 infraspecific taxa of C. angustifolia described from Lāna'i and Maui. These plants form a morphologically cohesive taxon that is clearly distinguishable from C. angustifolia and worthy of specific rank (Lammers, 1988b).

13. Cyanea fauriei H. Lév.

[C. coriacea (A. Gray) Hillebr. var. degeneriana F. Wimmer; C. c. var. fauriei (H. Lév.) F. Wimmer; C. c. f. gratiosa F. Wimmer; Delissea coriacea A. Gray var. degeneriana (F. Wimmer) St. John; D. c. var. gratiosa (F. Wimmer) St. John; D. fauriei (H. Lév.) St. John, nom. illeg.]

(end)

Branched or unbranched shrubs 2.5-4 m tall. Leaves narrowly oblong to oblong, blades 17-30 cm long, 4-9 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or very sparsely pubescent along the midrib, margins callose-crenulate, apex acute to obtuse, base obtuse to cuneate or attenuate, petioles 5-16 cm long. Inflorescences 12-25-flowered, pendent, peduncles 60-240 mm long, rachis 30-120 mm long, pedicels 12-20 mm long; hypathium obovoid, 3-5 mm long, 2-4 mm wide; calyx lobes dentiform, 0.5-1 mm long, apex acute to acuminate; corolla white striped or tinged purplish, 20-30 mm long, 2-3 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 8-10 mm long. Occurring in mesic valleys, 300-600 m, eastern and southern Kaua'i. — Plate 2.

page 449

When St. John (1987c) made the new combination Delissea fauriei (H. Lév.) St. John, he apparently overlooked an earlier homonym, Delissea fauriei (H. Léveillé, 1913b). The type of the latter (Faurie 572) could not be located and it is regarded as species dubia (Wimmer, 1943).

14. Cyanea fissa (H. Mann) Hillebr.

[Delissea fissa H. Mann; Cyanea fissa subsp. gayana (Rock) Lammers; C. gayana Rock; C. g. var. duvelii Rock; C. g. var. wainihaensis Rock; C. humilis Wawra; C. multispicata H. Lév.; Delissea gayana (Rock) St. John; D. g. var. duvelii (Rock) St. John; D. g. var. wainihaensis (Rock) St. John; D. humilis (Wawra) St. John; D. multispicata (H. Lév.) St. John]

(end)

Unbranched or sparingly branched shrubs 1-5 m tall, with yellow latex. Leaves oblanceolate to obovate, blades 32-80 cm long, 5-13 cm wide, upper surface green, glabrous, lower surface pale green, pubescent, margins minutely callose-denticulate, apex acuminate to acute, base attenuate, narrowing imperceptibly into a winged petiole 0.5-4 cm long. Inflorescences 8-20-flowered, pubescent, peduncles 20-60 mm long, pedicels 8-20 mm long; hypanthium obconical to obovoid, 6-10 mm long, 4-6 mm wide, pubescent; calyx lobes triangular to narrowly triangular, 3-16 mm long, 1.5-2.5 mm wide, apex acute to acuminate; corolla white to pale lilac, longitudinally striped with purple, 25-35 mm long, 4-5 mm wide, densely pubescent, the tube gently curved, the lobes spreading; staminal column glabrous; anthers minutely papillose, the lower 2 with apical tufts of white hairs. Berries reddish orange, broadly obovoid, 13-20 mm long, 10-15 mm wide. Occurring in bogs, diverse mesic forest, and wet forest, 200-915 m, Kaua'i. — Plate 53.

Two subspecies may be distinguished: subsp. fissa [incl. Cyanea gayana var. wainihaensis, C. humilis] with calyx lobes 6-16 mm long (Wainiha to Keālia, 200-300 m, northern Kaua'i); and subsp. gayana [incl. C. gayana var. duvelii, C. multispicata] with calyx lobes 3-5 mm long (Koai'e Stream to Hanahanapuni and Hā'upu Ridge, 300-915 m, southern Kaua'i). The specimen selected by St. John (1971a) as the lectotype of C. multispicata (Faurie 594, BM, G, P) is a specimen of C. fissa subsp. gayana. If that taxon is recognized as a species, the name Cyanea multispicata has priority over C. gayana.

15. Cyanea giffardii Rock

[Delissea giffardii (Rock) St. John]

(end, Ex)

Palm-like trees 5-10 m tall, with yellow latex. Leaves oblanceolate, blades 50-60 cm long, 12-15 cm wide, upper surface glabrous, lower surface pubescent, margins undulate, minutely callose-denticulate, shallowly lobed at base, apex acuminate, base cuneate, petioles 3.5-6.5 cm long. Inflorescences 5-10-flowered, pendent, hanging below the leaves, peduncles 100-180 mm long, pedicels 2-3 mm long; hypanthium obconical, 16-18 mm long, 14-15 mm wide, glabrous; calyx lobes deltate, ca. 5 mm long, ca. 5 mm wide, apex acute; corolla blackish purple, 70-80 mm long, 5-8 mm wide, glabrous, the tube curved, the lobes 3/4 as long to as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries blackish purple, globose, ca. 25 mm long. Known only from the type material [Rock 12802 (coll Giffard), BISH, CAS; Rock & Holm 12802b, BISH, L, NSW, NY], collected in 1917 in wet forest, 725-800 m, near Glenwood, Hawai'i.

16. Cyanea glabra (F. Wimmer) St. John

[C. knudsenii Rock var. glabra F. Wimmer; C. holophylla Hillebr. var. obovata Rock; C. scabra Hillebr. var. variabilis Rock, pro parte; Delissea glabra (F. Wimmer) St. John; D. holophylla (Hillebr.)St. John var. obovata (Rock) St. John; D. scabra (Hillebr.) St. John var. variabilis (Rock) St. John]

(end, Ex)

Branched shrubs, height unknown. Leaves dimorphic: juveniles pinnately cleft, parted, or divided, blades 20-46 cm long, 10-20 cm wide (across the segments), upper surface green, glabrous, lower surface pale green, glabrous or sparsely pubescent, both surfaces often aculeate on the midrib and veins, apex acuminate, base truncate or obtuse, segments 5-8 per side, margins callose-denticulate to callose-dentate, petioles 9-15 cm long, aculeate; adults elliptic to oblanceolate, blades 23-36 cm long, 7-12 cm wide, upper surface green, glabrous,

page 450

page 451

lower surface pale green, glabrous or sparsely pubescent, margins callose-denticulate to irregularly lobed, apex acuminate, base obtuse to cuneate, petioles 4-11 cm long, smooth or sparsely aculeate. Inflorescences 6-8-flowered, peduncles 20-55 mm long, pedicels 12-25 mm long; hypanthium obconical, 7-10 mm long, 4-5 mm wide; calyx lobes triangular or narrowly oblong, 2-8 mm long, 1-2 mm wide, apex acute to acuminate; corolla white, often tinged pale lilac, 50-60 mm long, 8-9 mm wide, glabrous, the tube curved, the lobes spreading, 1/4-1/3 as long as the tube, sparsely muricate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellowish orange, ellipsoid, 10-15 mm long, crowned by the persistent calyx lobes. Formerly occurring in wet forest, windward slopes of Haleakalā, East Maui. Last collected in 1920 (Forbes 2598.M, BISH).

This species is not at all related to Cyanea knudsenii of Kaua'i (here included in C. sylvestris), but rather to C. longissima of East Maui, C. platyphylla of Hawai'i, and C. scabra of West Maui. Cyanea scabra var. variabilis is here interpreted to comprise 2 distinct elements. The holotype is Cyanea longissima, while the paratypes are C. glabra.

17. Cyanea grimesiana Gaud.

[C. g. subsp. cylindrocalyx (Rock) Lammers; C. g. var. cylindrocalyx Rock; C. g. var. hirsutifolia Rock; C. g. var.lydgatei Rock; C. g. var. mauiensis Rock; C. g. var. munroi Hosaka; C. g. subsp. obatae (St. John) Lammers; C. g. var.obatae St. John; C. lobata H. Mann var. hamakuae Rock; Delissea grimesiana (Gaud.) St. John; D. g. var. cylindrocalyx (Rock) St. John; D. g. var. hirsutifolia (Rock) St. John; D. g. var. lydgatei (Rock) St. John; D. g. var. mauiensis (Rock) St. John; D. g. var. munroi (Hosaka) St. John; D. g. var. obatae (St. John) St. John; D. lobata (H. Mann) St. John var. hamakuae (Rock) St. John; Lobelia grimesiana (Gaud.) Hook. & Arnott]

(end, [1 subsp, E*])

Shrubs 1-3.2 m tall; stems unbranched or sparingly branched, muricate. Leaves pinnately divided, blades 27-58 cm long, 14-32 cm wide (across the segments), upper surface glabrous, rarely sparsely pubescent or muricate, lower surface glabrous or muricate along the midrib, rarely densely pubescent, segments 9-12 per side, oblong, lanceolate, narrowly triangular, or narrowly ovate, 0.8-5 cm wide, apex acuminate, apiculate, margins or segments irregularly lobed to irregularly cleft, the lobes acute to obtuse, apiculate, petioles 8-32 cm long, muricate. Inflorescences 6-12-flowered, peduncles 15-75 mm long, pedicels 20-50 mm long, bracts linear to lanceolate or narrowly oblong, 12-35 mm long, 1-3 mm wide; hypanthium obconical, 7-16 mm long, 3-8 mm wide; calyx lobes ovate to lanceolate, overlapping at base, 10-44 mm long, 4-14 mm wide, or linear to narrowly triangular, separated at base by narrow sinuses, 3-13 mm long, 1-2 mm wide, or rarely connate 7-25 mm from the base, forming an irregularly 5-lobed tube 20-30 mm long, 8-12 mm wide; corolla purplish or greenish to yellowish white, often suffused or striped with magenta, 55-80 mm long, 5-10 mm wide, glabrous or very sparsely pubescent, rarely densely pubescent, the tube arcuate, the lobes spreading, 1/4-1/2 as long as the tube; staminal column glabrous; anthers glabrous or sometimes pubescent at base, all 5 (very rarely only the lower 2) with apical tufts of white hairs. Berries orange, obovoid to ellipsoid, 18-30 mm long, 8-15 mm wide, crowned by the persistent calyx lobes. Occurring in mesic to wet forest, 200-730 m, on O'ahu, Moloka'i, Lāna'i, Maui, and Hawai'i. — Plate 53.

Three subspecies may be distinguished: subsp. cylindrocalyx with calyx lobes connate 7-25 mm from the base, forming an irregularly 5-lobed tube 20-30 mm long, 8-12 mm wide, known only from 2 specimens (Rémy 309, P; and Rock 4629, BISH), collected in 1851-1855 and 1909, respectively, at Waipi'o Valley, Hawai'i; subsp. grimesiana [incl. Cyanea grimesiana var. lydgatei, C. g. var. mauiensis, C. g. var. munroi, C. lobata var. hamakuae] with calyx lobes ovate to lanceolate, 10-44 mm long, 4-14 mm wide, overlapping at the base, which occurs primarily in the Ko'olau Mountains, O'ahu, but also on Mount Ka'ala in the northern Wai'anae Mountains, O'ahu, and on Moloka'i, Lāna'i, and Maui; and subsp. obatae [incl. C. grimesiana var. hirsutifolia] with calyx lobes linear to narrowly triangular, 3-13 mm long, 1-2 mm wide, separated at the base by narrow sinuses, from

page 452

Pu'uhāpapa to Ka'aikukui, southern Wai'anae Mountains, O'ahu. Gaudichaud (1829a) gives "Toué-noui" [kuenui] as the Hawaiian name for the species.

18. Cyanea hamatiflora Rock

[C. carlsonii Rock; C. hamatiflora subsp. carlsonii (Rock) Lammers; Delissea carlsonii (Rock) St. John; D. hamatiflora (Rock) St. John]

(end, [1 subsp. E*])

Palm-like trees, 3-8 m tall, with tan latex. Leaves oblanceolate to narrowly oblong, blades 50-80 cm long, 8-14 cm wide, upper surface glabrous or sparsely pubescent, lower surface pubescent at least along the midrib and veins, margins minutely callose-denticulate, apex acute, sessile. Inflorescences 5-10-flowered, peduncles 15-30 mm long, pedicels 5-12 mm long; hypanthium obconical to obovoid, 12-30 mm long, 6-12 mm wide; calyx lobes narrowly oblong, 12-45 mm long; 3-6 mm wide, apex obtuse and apiculate; corolla magenta, 60-80 mm long, 6-11 mm wide, glabrous, the tube suberect to slightly curved, the lobes 1/4-1/2 as long as the tube, all curved downward in the same plane, the corolla thus appearing unilabiate; staminal column glabrous or pubescent at base; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purplish red, ellipsoid to obovoid, 30-45 mm long, 20-27 mm wide, crowned by the persistent calyx lobes. Occurring in wet forest, 1,200-1,740 m, East Maui and Hawai'i.

Two subspecies may be distinguished: subsp. carlsonii with calyx lobes 30-45 mm long, 4-6 mm wide, and pedicels 8-12 mm long (rare, Kona District, 1,300-1,740 m, Hawai'i); and subsp. hamatiflora with calyx lobes 12-18 mm long, 3-4 mm wide, and pedicels 5-6 mm long (windward slopes of Haleakalā, 1,200-1,400 m, East Maui).

19. Cyanea hardyi Rock

[C. coriacea (A. Gray) Hillebr. var. hardyi (Rock) F. Wimmer; Delissea coriacea A. Gray var. hardyi (Rock) St. John]

(end)

Branched shrubs or trees 1-7 m tall. Leaves elliptic to oblanceolate, blades 18-28 cm long, 2.5-5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or sparsely pubescent along the midrib, margins callose-crenulate, apex acute to acuminate, base attenuate into a winged petiole 2.5-8 cm long. Inflorescences 15-30-flowered, pendent, peduncles 60-240 mm long, rachis 20-100 mm long, pedicels 8-12 mm long; hypanthium obconical, 3-5 mm long, 2-4 mm wide; calyx lobes dentiform, 0.5-1 mm long, apex acute to acuminate; corolla purplish to blackish purple, 20-26 mm long, 2-3 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 8-10 mm long. Occurring in mesic valleys and diverse mesic forest, 425-790 m, southern Kaua'i.

20. Cyanea hirtella (H. Mann) Hillebr.

[Delissea hirtella H. Mann; Cyanea chockii Rock; C. communis Rock; C. hirtella var. striata F. Wimmer; C. h. var. subglabra F. Wimmer; Delissea chockii (Rock) St. John; D. hirtella var. striata (F. Wimmer) St. John; D. h. var. subglabra (F. Wimmer) St. John]

(end)

Unbranched or sparingly branched shrubs 1-6 m tall. Leaves oblanceolate to oblong, blades 10-27 cm long, 2-6 cm wide, upper surface green, glabrous or sparsely pubescent, lower surface pubescent to subglabrous, margins minutely callose-denticulate to callose-serrulate, apex acuminate, base attenuate to obtuse, petioles 2.5-6 cm long. Inflorescences 5-12-flowered, pubescent, peduncles 10-40 mm long, pedicels 6-15 mm long; hypanthium obconical to obovoid, 6-11 mm long, 4-5 mm wide, pubescent; calyx lobes narrowly triangular, 1-3 mm long, 0.5-1 mm wide, apex acuminate; corolla white to pale purplish with purple longitudinal stripes, 30-40 mm long, 3-4 mm wide, densely pubescent, the tube suberect, the lobes spreading; staminal column glabrous; anthers minutely and sparsely pubescent, the lower 2 with apical tufts of white ahris. Berries orange, globose to obovoid, 10-16 mm long, 6-9 mm wide. Occurring in wet forest, 1,030-1,400 m, western Kaua'i. — Plate 53.

page 453

21. Cyanea horrida (Rock) Degener & Hosaka

[C. ferox Hillebr. var. horrida Rock; Delissea horrida (Rock) St. John]

(end) Hāhā nui

Palm-like trees 1-4 m tall. leaves pinnately parted or divided more so in juveniles, blades 20-37 cm long, 10-19 cm wide (across the segments), upper surface green, muricate (aculeate in juveniles), lower surface pale green, pubescent and muricate (aculeate in juveniles), segments 8-12 per side, narrowly oblong, 15-35 mm wide, apex acute, margins irregularly dentate or lobed (cleft, parted, or divided in juveniles), the lobes acute to obtuse, petioles 4-13 cm long, muricate (aculeate in juveniles). Inflorescences 5-8-flowered, clustered among the leaves, muricate, peduncles 20-60 mm long, pedicels 15-25 mm long, bracts linear, 15-30 mm long, 1-2 mm wide; hypanthium oblong to broadly ellipsoid, 10-15 mm long, 5-10 mm wide; calyx lobes narrowly triangular, 8-24 mm long, 2-4 mm wide; corolla blackish purple or greenish white suffused with purple, 70-80 mm long, 7-10 mm wide, densely pubescent to muricate externally, glabrous within, the tube arcuate, the lobes spreading, 1/2-3/4 as long as the tube; staminal column glabrous; anthers glabrous, all 5 or only the lower 2 with apical tufts of white hairs. Berries yellow, ellipsoid, 14-25 mm long, crowned by the persistent calyx lobes. Occurring in wet forest and margins of subalpine forest, 1,520-2,000 m, windward slopes of Haleakalā, East Maui.

Several specimens appear to represent hybrids between Cyanea horrida and C. macrostegia subsp. macrostegia, including the type of C. atra Hillebr. var. lobata Rock (Rock 8637, A, BISH, CAS, GH, NSW, US).

22. Cyanea kunthiana Hillebr.

[C. bishopii Rock, nom. illeg.; Delissea bishopii (Rock) St. John, nom. illeg.]

(end)

Shrubs 0.5-1.5 m tall; stems subherbaceous, woody only at base, unbranched or sparingly branched. Leaves elliptic to narrowly obovate, blades 12-30 cm long, 3-7 cm wide, upper surface green, sparsely pubescent, lower surface whitish green, pubescent, margins minutely callose-denticulate, apex acuminate, base cuneate, petioles 2-4 cm long. Inflorescences 8-24-flowered, pubescent, peduncles 6-25 mm long, pedicels 6-15 mm long; hypanthium obovoid, 4-6 mm long, pubescent; calyx lobes narrowly triangular, apex acuminate, 5-8 mm long; corolla whitish with pale lilac longitudinal stripes or dark purplish, 30-35 mm long, 3-4 mm wide, sparsely pubescent, the tube suberect to slighly curved, the lobes spreading, 1/4-1/3 as long as the tube; staminal column sparsely pubescent; anthers densely pubescent, the lower 2 with apical tufts of white hairs. Berries purplish orange, obovoid to ellipsoid, 7-8 mm long. Occurring in wet forest, 1,000-1,600 m, Maui. — Plate 53.

Cyanea kunthiana is interpreted here as a new taxon described by Hillebrand (1888) on the basis of a new type, rather than as a new combination based on Delissea kunthiana Gaud. Hillebrand's discussion makes it quite clear that he did not accept the identity of his plant and Gaudichaud's. Delissea kunthiana, while validly published, was accompanied only by a plate and lacked any descriptive text. Because the type cannot be located and the plant is not identifiable from the plate, D. kunthiana must remain species incognita. Cyanea bishopii, based on the same type as C. kunthiana, is superfluous.

23. Cyanea leptostegia A. Gray

[C. l. var. velutina Skottsb.; Delissea coriacea A. Gray var. pinnatiloba A. Gray ex H. Mann; D. leptostegia (A. Gray) St. John; D. l. var. velutina (Skottsb.) St. John]

(end) Hāhā lua

Palm-like trees 3-14 m tall, with bright yellow latex. Leaves dimorphic: juveniles irregularly parted or divided, blades 20-70 cm long, 5-12 cm wide (across the segments), apex acute to rounded, segments 4-7 per side, margins minutely callose-denticulate, petioles 0.5-2 cm long; adults linear to narrowly oblanceolate, 50-100 cm long, 2.5-8.5 cm wide, glabrous or lower surface

page 454

rarely pubescent, margins gently undulate and minutely callose-denticulate, apex acuminate, sessile. Inflorescences 10-25 flowered, horizontal, densely clustered among the leaves, peduncles 20-80 mm long, pedicels 10-20 mm long; hypanthium oblong, 15-25 mm long; calyx lobes linear, 30-55 mm long, 0.5-1 mm wide; corolla rose to purple, 45-55 mm long, 3-5 mm wide, glabrous, the tube suberect, all 5 lobes curved downward in the same plane, the corolla thus appearing unilabiate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellow, ovoid, 20-35 mm long, 8-12 mm wide. [2n = 28*.] Occurring in diverse mesic forest, 1,000-1,300 m, western Kaua'i. — Plate 54.

Cyanea leptostegia does not flower precociously. However, leaves of juveniles are described because they are commonly encountered, radically different in appearance from those of adults, and were the basis for the description of Delissea coriacea var. pinnatiloba.

24. Cyanea linearifolia Rock

[Delissea linearifolia (Rock) St. John]

(end, Ex)

Unbranched shrubs 2-2.5 m tall. Leaves linear, 28-33 cm long, 0.3-1 cm wide, margins revolute, sessile. Inflorescences 9-12-flowered, glabrous, peduncles 45-70 m long, pedicels 12-15 mm long; calyx lobes dentiform, 0.5-1 mm long, apex acute to acuminate; corolla 15-20 mm long, 1.5-2 mm wide, glabrous, the lobes spreading. Known only from the holotype (Lydgate s.n., BISH), collected in the Hi'i [Wahiawa] Mountains of Kaua'i, probably in wet forest, and 3 additional specimens lacking any collection data (BISH; possible isotype).

25. Cyanea lobata H. Mann

[C. baldwinii C. Forbes & G. Munro; Delissea baldwinii (C. Forbes & G. Munro) St. John; D. lobata (H. Mann) St. John]

(end, E*)

Shrubs 1.3-2.3 m tall; stems sparingly branched, smooth or sparsely muricate. Leaves oblong to obovate, blades 30-50 cm long, 10-15 cm wide (across the lobes), upper surface glabrous, lower surface sparsely pubescent or muricate along the midrib and veins, apex acuminate, margins irregularly lobed, the lobes 12-25 per side, deltate to triangular, petioles 7-22 cm long, sparsely muricate. Inflorescences 5-12-flowered, peduncles 30-75 mm long, pedicels 18-35 mm long; hypanthium obconical, 8-12 mm long, 3-6 mm wide; calyx lobes oblong, 10-24 mm long, 3-7 mm wide, apex obtuse, apiculate; corolla greenish white or purplish, 60-70 mm long, 5-11 mm wide, pubescent at least on the lobes, the tube arcuate, the lobes spreading, 1/4-1/2 as long as the tube; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Berries yellow, globose. Rare in mesic valleys, 550-900 m, Lāna'i and northern West Maui.

This species, which had not been collected on Maui since Hillebrand's time, was rediscovered at Waikapū Valley in 1982 (Hobdy 1675, BISH). On Lāna'i, a single individual was discovered at Pu'uali'i in 1919 (Munro 674, BISH, NY; the type of C. baldwinii); it was still alive in 1934 (Degener, 1936). This individual was propagated by Munro and the young plants were set out at Lāna'ihale and Waikeakua, Lāna'i, and at Munro's home on O'ahu. Plants persisted at the first site until at least 1935 (Fosberg 12147, POM) and at Munro's home until at least 1940 (Munro s.n., K). Cyanea lobata var. hamakuae is treated here as a synonym of C. grimesiana subsp. grimesiana.

26. Cyanea longissima (Rock) St. John

[C. scabra Hillebr. var. longissima Rock; C. s. var variabilis Rock, pro parte; Delissea longissima (Rock) St. John]

(end,Ex)

Habit unknown. Leaves oblanceolate to obovate, blades 27-45 cm long, 4.5-11 cm wide, upper surface green, glabrous, lower surface whitish green, glabrous, margins remotely callose-denticulate, apex acute to acuminate, base long-attenuate, narrowing imperceptibly into a winged petiole 4-7 cm long. Inflorescenses 8-10-flowered, peduncles 10-15 mm long, pedicels 5-12 mm long; hypanthium obconical, 4-6 mm long,

page 455

page 456

3-4 mm wide; calyx lobes narrowly triangular, 3-4 mm long, 1-2 mm wide, apex acuminate; corolla white, sometimes with pale lilac longitudinal stripes, 50-60 mm long, 5-9 mm wide, glabrous, the tube arcuate, the lobes spreading, 1/4-1/3 as long as the tube, muricate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries orange, obovoid to ellipsoid, 10-12 mm long. Formerly occurring in wet forest, windward slopes of Haleakalā, East Maui. Last collected in 1927 (Degener 18071, A, MO, NY).

As here interpreted, Cyanea scabra var. variabilis comprises 2 distinct elements. The holotype [Rock 10259 (coll. Lyon), BISH] is referable to the present species, while the paratypes are referred to Cyanea glabra.

27. Cyanea macrostegia Hillebr.

[C. atra Hillebr.; C. bicolor St. John; C. gibsonii Hillebr.; C. hanaensis St. John; C. macrostegia subsp. gibsonii (Hillebr.) Lammers; C. m. var. parvibracteata Rock; C. mariana F. Wimmer; Delissea atra (Hillebr.) St. John; D. bicolor (St. John) St. John; D. gibsonii (Hillebr.) St. John; D. macrostegia (Hillebr.) St. John; D. m. var. parvibracteata (Rock) St. John; D. mariana (F. Wimmer) St. John]

(end [1 subsp. E*]

Palm-like trees 1-7 m tall. Leaves oblanceolate to oblong or elliptic, blades 20-80 cm long, 6.5-20 cm wide, upper surface glabrous or sparsely pubescent, lower surface pubescent, margins callose-denticulate, rarely erose-dentate, apex acute to obtuse, rarely acuminate, base cuneate, obtuse, or rounded, petioles 3-17 cm long, often muricate. Inflorescences 5-15-flowered, clustered among the leaves, peduncles 10-80 mm long, pedicels 10-25 mm long; hypanthium obconical, 10-20 mm long, 5-10 mm wide; calyx lobes oblong, narrowly oblong, or ovate,6-55 mm long, 4-10 mm wide, apex obtuse, truncate, or emarginate, rarely acute, often with an apiculation; corolla blackish purple externally, white or pale lilac within, 60-80 mm long, 6-11 mm wide, densely pubescent or rarely subglabrous externally, glabrous within, the tube suberect or arcuate, the lobes spreading, 1/2-3/4 as long as the tube; staminal column glabrous or sparesely pubescent; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellowish orange, obovoid or ellipsoid, 15-30 mm long, 12-25 mm wide, crowned by the persistent calyx lobes. Occurring in wet forest, 760-1,970 m, Lāna'i and Maui. — Plate 54.

Two subspecies may be distinguished: subsp. gibsonii with the corolla tube arcuate (rare, Lāna'i); and subsp macrostegia [incl. Cyanea atra, C. bicolor, C. hanaensis, C. macrostegia var. parvibracteata, C. mariana] with the corolla tube suberect (East and West Maui). Size and shape of the calyx lobes and pubescence of the corolla vary markedly in the latter, but it is not possible to distinguish additional meaningful subspecies on this basis. Although the Maui race is still common, the Lāna'i race has been collected only twice since 1927, in 1971 (Hobdy 235, BISH) and in 1984 (Hobdy 2197, BISH). Cyanea atra var lobata appears to represent hybrids of C. horrida and C. macrostegia subsp. macrostegia, while C. macrostegia var. viscosa may represent hybrids between C. aculeatiflora and C. macrostegia subsp. macrostegia.

28. Cyanea mannii (Brigham) Hillebr.

[Delissea mannii Brigham]

(end,E*)

Branched shrubs 1.5-3 m tall. Leaves narrowly elliptic to lanceolate, blades 12-21 cm long, 2.5-5 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent, margins callose-serrulate, apex acuminate, base obtuse to cuneate, petioles 2.2-10 cm long. Inflorescences 6-12-flowered, peduncles 20-35 mm long, pedicels 8-12 mm long; hypanthium broadly obconical, 4-6 mm long, 3-5 mm wide; calyx lobes triangular, 3-5 mm long, 2-3 mm wide, apex acute; corolla purplish, 30-35 mm long, 3-4 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Occurring in mesic to wet forest, 1,000-1,080 m, central Moloka'i. — Plate 54.

This species, which had not been collected since 1912 (Forbes 33.Mo, A;

page 457

Forbes 265.Mo, A, BH, BISH, CAS, DS, K, L, MO, NY, P, US, W, WELT), was rediscovered in 1984 (Lammers et al. 5703, OS; Lammers et al. 57909, BISH, OS; Misaki s.n., OS).

29. Cyanea marksii Rock

[C. tritomantha A. Gray var. lydgatei Hillebr. ex Rock; Delissea marksii (Rock) St. John; D. tritomantha (A. Gray) St. John var. lydgatei (Hillebr ex Rock) St. John]

(end)

Palm-like trees 2 m tall. Leaves dimorphic: juveniles obovate to oblong, blades 13-18 cm long, 5.5-7 cm wide, upper surface aculeate, lower surface pubescent and aculeate, margins callose-denticulate to callose-dentate, apex obtuse to acute, base rounded to obtuse, petioles 2.5-4 cm long, aculeate; adults oblong to oblanceolate, blades 25-31 cm long, 8-12 cm wide, upper surface smooth or muricate, lower surface pubescent, margins minutely callose-denticulate, apex acute, base cuneate, petioles 8-14 cm long, sparsely muricate. Inflorescences 5-8-flowered, clustered among the leaves, peduncles 10-20 mm long, pedicels 10-20 mm long; hypanthium oblong, 12-15 mm long, 7-3 mm wide, pubescent; calyx lobes narrowly triangular, 8-10 mm long, 2-4 mm wide, apex acute; corolla blackish purple externally, white within, 70-80 mm long, 4-7 mm wide, pubescent, the tube arcuate, the lobes spreading, 1/2-3/4 as long as the tube, sparsely muricate; staminal column glabrous to sparsely pubescent; anthers glabrous or pubescent, the lower 2 with apical tufts of white hairs. Berries ellipsoid, 14-18 mm long, crowned by the persistent calyx lobes. Occurring in wet forest, 925-1,200 m, Kona District, Hawai'i.

In vegetative features, the type (Rock 25631, BISH) and subsequent collections of Cyanea marksii resemble juvenile individuals of certain palm-like species (e.g., C. macrostegia, C. tritomantha). Further, C. marksii bears large, blackish purple flowers similar to those found in adults of those species. On this bases, Carlquist (1974, 1980) suggested that C. marskii might be a "permanently juvenile" or paedomorphic species, one that becomes capable of reproduction but never attains adult morphology. However, the adult plant has now been recognized. Cyanea tritomantha var.lydgatei is a palm-like plant known only from the holotype (Lydgate s.n., BISH), collected in 1873 on the Kona coast, Hawai'i. It differs from C. tritomantha in its shorter calyx lobes, longer corolla, and shorter peduncles, but it is identical to C. marksii in these features. Accordingly, C. marksii is interpreted as a palm-like species related to C. macrostegia and C. tritomantha, which is known primarily from precociously flowering juvenile individuals. A careful search of the Kona forests for adult Cyanea marksii is needed to confirm this interpretation.

30. Cyanea mceldowneyi Rock

[Delissea mceldowneyi (Rock) St. John]

(end,E*)

Shrubs 2-3 m tall; stems unbranched, muricate to aculeate. Leaves dimorphic: juveniles broadly elliptic to obovate, blades 15-22 cm long, 5-9 cm wide, upper surface green, aculeate, lower surface pale green, pubescent and aculeate, margins callose-dentate, apex acuminate, base rounded, petioles 2.5-4.5 cm long, aculeate; adults oblanceolate to narrowly oblong, blades 20-35 cm long, 5-9 cm wide, upper surface green, glabrous or sparsely muricate, lower surface pale green, very sparsely pubescent, margins callose-denticulate, apex acuminate, base cuneate, petioles 3.5-6 cm long, smooth or muricate. Inflorescences 5-7-flowered, peduncles 15-30 mm long, pedicels 10-14 mm long; hypanthium obconical, ca. 5 mm long, ca. 4 m wide; calyx lobes oblong, ca. 5 mm long, ca. 2.5 mm wide, apex emarginate and apiculate; corolla white with purple longitudinal stripes, ca. 40 mm long, ca. 8 mm wide, glabrous, the tube arcuate, the lobes spreading ca. 1/4 as long as the tube, muricate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Occurring in wet forest, 925-1,280 m, windward slopes of Haleakalā, East Maui

31. Cyanea membranacea Rock

[C. angustifolia (Cham.) Hillebr. var. tomentella Hillebr.; C. a. var. t. f. subpubescens F. Wimmer]

page 458

C. coriacea (A. Gray) Hillebr. var. serratifolia Rock; C. dentata F. Wimmer; Delissea angustifolia (Cham.) K. Presl var. tomentella (Hillebr.) St. John; D. a. var. t. f. subpubescens (F. Wimmer) St. John; D. coriacea A. Gray var. serratifolia (Rock) St. John; D. dentata (F. Wimmer) St. John; D. serratifolia (Rock) St. John]

(end)

Sparingly branched shrubs 2-4 m tall. Leaves oblanceolate to oblong, blades 18-32 cm long, 3-9 cm wide, upper surface green, glabrous or sparsely pubescent, lower surface pale green, pubescent, margins callose-serrate, apex obtuse or acute, base obtuse to cuneate, petioles 4-12 cm long. Inflorescences 7-12-flowered, peduncles 20-40 mm long, pedicels 5-20 mm long; hypanthium purple, broadly obovoid, 4-5 mm long, 3-4 mm wide; calyx lobes dentiform, 0.5-1 mm long, apex acute to acuminate; corolla white, sometimes slightly tinged pale purplish, 30-40 mm long, 3-4 mm wide, sparsely pubescent or rarely subglabrous, the tube suberect to gently curved, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 8-10 mm long. Occurring on shaded slopes and gulches in mesic forest, 580-730 m, Wai'anae Mountains, O'ahu.

32. Cyanea obtusa (A. Gray) Hillebr.

[Delissea obtusa A. Gray]

(end,Ex)

Shrubs 2-5 m tall; stems branched, pubescent. Leaves oblong to oblanceolate, blades 15-30 cm long, 4-9 cm wide, upper surface green, sparsely pubescent, lower surface pale green, densely pubescent, margins coarsely callose-dentate, apex acute, base cuneate, petioles 3.5-10 cm long. Inflorescences 6-12-flowered, densely pubescent, peduncles 20-50 mm long, pedicels 15-25 mm long; hypanthium obovoid, 4-6 mm long, densely pubescent; calyx lobes narrowly triangular, 1-2 mm long, densely pubescent throughout, apex acute to acuminate; corolla purplish, 30-36 mm long, 1.5-3 mm wide, densely pubescent, the tube 1.5-3 mm wide, densely pubescent, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Formerly occurring in wet forest. Maui. Last collected in 1919 (Forbes 1290.M, BISH, S, W).

Two specimens (Rock 8517, BISH, CAS, NSW; Faurie 563, A, P, G, W) appear to represent hybrids between Cyanea obtusa and C. elliptica. Delissea obtusa var. mollis from Mauna Kea, Hawai'i, treated as a synonym of Cyanea obtusa by Rock (1919a) and others, is here regarded as a synonym of Clermontia pyrularia.

33. Cyanea pilosa A. Gray

[C. longipedunculata Rock; C. megacarpa (Rock) Rock; C. pilosa subsp. longipedunculata (Rock) Lammers; C. p. var. megacarpa Rock; Delissea longipedunculata (Rock) St. John; D. megacarpa (Rock) St. John; D. pilosa (A. Gray) H. Mann]

(end)

Shrubs 0.8-2 m tall; stems subherbaceous, woody only at base, unbranched or rarely sparingly branched from base, pubescent. Leaves broadly elliptic to broadly obovate, blades 15-42 cm long, 8-15 cm wide, upper surface green, pubescent, lower surface whitish green, densely pubescent, margins erose, apex acuminate, base rounded to cuneate, petioles 3-9 cm long. Inflorescences 6-28-flowered, pubescent, peduncles 15-110 mm long, rachis 5-70 mm long, pedicels 5-10 mm long; hypanthium obovoid, 4-6 mm long; calyx lobes narrowly triangular, 2-5 mm long, apex acuminate; corolla white tinged or streaked with purple, 20-28 mm long, 3-4 mm wide, glabrous, the tube suberect to gently curved, the lobes spreading, 1/4-1/3 as long as the tube; staminal column white, pubescent toward apex; anthers white, pubescent, the lower 2 with apical tufts of white hairs. Berries orange, globose to ellipsoid, 7-22 mm long, crowned by the persistent calyx lobes. Occurring in wet forest, 610-1,520 m, Kohala Mountains, windward Mauna Kea, and windward Mauna Loa, Hawai'i.

Two subspecies may be distinguished: subsp. longipedunculata with 12-28-flowered inflorescences 70-180 mm long and berries 7-10 mm long (windward Mauna Kea and Mauna Loa, 610-1,300m); and subsp. pilosa [incl. Cyanea pilosa var. megacarpa] with 6-12-flowered inflorescences 20-70 mm long and berries 10-22 mm long (Kohala Mountains and windward Mauna Kea, 1,000-1,520m).

page 459

34. Cyanea pinnatifida (Cham.) F. Wimmer

[Lobelia pinnatifida Cham.; Cyanea selachicauda Degener; Delissea pinnatifida (Cham.) K. Presl; Rollandia pinnatifida (Cham.) G. Don]

(end, E*)

Shrubs 0.8-3 m tall; stems unbranched or sparingly branched, glabrous or sparsely muricate. Leaves pinnately parted, cut 1/2-3/4 the distance to the midrib, blades 25-60 cm long, 3.5-8 cm wide (between the segments), 8-25 cm wide (across the segments), upper surface glabrous, lower surface sparsely pubescent, segments 2-6 per side, narrowly oblong, 1-2.5 cm wide, margins minutely callose-denticulate, apex acute or obtuse, petioles 3-7 cm long. Inflorescences 8-15-flowered, pubescent, peduncles 40-50 mm long, pedicels 10-15 mm long; hypanthium obconical, 10-12 mm long, 6-8 mm wide, pubescent; calyx lobes triangular, 3-5 mm long, 2-2.5 mm wide, apex obtuse; corolla greenish white with purplish longitudinal stripes at base, 36-50 mm long, 4-5 mm wide, sparsely pubescent, the tube suberect to slightly curved, the lobes spreading; staminal column white, sparsely pubescent at base; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Rare in diverse mesic forest, central Wai'anae Mountains, O'ahu. Last collected in 1965 (Carlquist & Parratt 1808, DUKE, MICH, RSA). — Plate 54.

Carlquist (1974, 1980) considered this species to be based upon precociously flowering juvenile individuals of another taxon found in the Wai'anae Mountains, Rollandia kaalae (here treated as a synonym of R. lanceolata subsp. calycina). However, these putative juveniles produce flowers in which the staminal column is not adnate to the corolla. In Rollandia, the staminal column is adnate to the corolla for 1/3 to 1/2 its length. Furthermore, in true juvenile individuals of Rollandia lanceolata, the leaves are not parted but rather are coarsely callose-dentate (e.g., Lammers & Buelow 5350, OS; Forbes 2568.O, BISH; Skottsberg 350, BISH, GB; and St. John 11038, BISH, US). Specimens purporting to show juvenile and adult foliage (Carlquist 1715, BISH, L, MICH, RSA) are mixed collections of Cyanea pinnatifida and Rollandia lanceolata subsp. calycina, respectively.

35. Cyanea platyphylla (A. Gray) Hillebr.

[Delissea platyphylla A. Gray; Cyanea bryanii Rock; C. crispohirta F. Wimmer; C. fernaldii Rock; C. nolimetangere Rock; C. pulchra Rock; C. rollandioides Rock; Delissea bryanii (Rock) St. John; D. crispohirta (F. Wimmer) St. John; D. fernaldii (Rock) St. John; D. nolimetangere (Rock) St. John; D. pulchra (Rock) St. John; D. rollandioides (Rock) St. John]

(end) 'Akū'akū

Shrubs 1-3 m tall; stems unbranched, smooth to muricate or aculeate (densely so in juveniles). Leaves dimorphic: juveniles oblong to obovate, blades 9-21 cm long, 4-7.5 cm wide, upper surface aculeate, lower surface pubescent and aculeate, margins callose-dentate to lobed, apex acute to acuminate, base rounded to obtuse, petioles 1.5-4 cm long, glabrous or pubescent, aculeate; adults oblong to oblanceolate or obovate, blades 28-68 cm long, 7-20 cm wide, upper surface glabrous or sparsely muricate to minutely aculeate, lower surface pale green, glabrous or sparsely pubescent to sparsely muricate, margins callose-denticulate to callose-dentate, apex acute to acuminate, base cuneate, petioles 6-19 cm long, glabrous or sparsely muricate. Inflorescences 6-25-flowered, peduncles 20-90 mm long, pedicels 10-25 mm long; hypanthium obconical, 5-9 mm long, 3-5 mm wide; calyx lobes narrowly oblong to narrowly triangular, 3-7 mm long, 1-2 mm wide, apex acuminate; corolla white, yellowish white with magenta stripes, or magenta, 42-54 mm long, 5-10 mm wide, glabrous or pubescent, the tube arcuate, the lobes spreading, 1/4-1/2 as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries pale orange, ellipsoid, 8-10 mm long, 6-8 mm wide. Occurring in wet forest, 120-915 m, Waipi'o Valley, windward Mauna Kea, Puna District, and windward Mauna Loa, Hawai'i.

Carlquist (1974, 1980) was correct when he indicated that Cyanea bryanii, C. fernaldii, C. nolimetangere, C. pulchra, and C. rollandioides represented different developmental stages of one species; C. crispohirta also belongs here. However, he erred in regarding Cyanea tritomantha as the fully adult stage. Cyanea platyphylla and C. tritomantha are not closely related. In the former, the corolla is 42-53 mm long, the tube arcuate, the lobes 1/4-1/2 as long as the tube; it is related to such species as

page 460

Cyanea scabra, C. glabra, C. longissima, and C. mceldowneyi. In contrast, Cyanea tritomantha has the corolla 60-75 mm long, the tube suberect to slightly curved, the lobes 1/2-3/4 as long as the tube; it is related to such species as C. aculeatiflora, C. macrostegia, and C. marksii. Carlquist apparently was misled by the remarkable similarity between juvenile individuals of Cyanea platyphylla and C. tritomantha, a resemblance noted earlier by Rock (1919a). However, while precocious flowering is apparently frequent in Cyanea platyphylla (e.g., Hatheway & Richards 539, F; Rock 10351, BISH), it has never been observed in C. tritomantha.

36. Cyanea pohaku Lammers

[Clermontia haleakalensis Rock, non Cyanea haleakalaensis St. John]

(end, Ex)

Sparingly branched trees 6-7 m tall. Leaves oblanceolate, blades 16-30 cm long, 1.5-4 cm wide, glabrous, margins callose-crenulate, apex subacute, base attenuate, narrowing imperceptibly into a winged petiole 0.8-2 cm long. Inflorescences 4-8-flowered, peduncles 18-48 mm long, pedicels 6-15 mm long, bracts narrowly triangular, 4-7 mm long; hypanthium obconical, 12-17 mm long; calyx lobes narrowly triangular, 4-7 mm long, 1-2 mm wide; corolla greenish white 30-45 mm long, 5-7 mm wide, the tube gently curved, the lobes 1/2-3/4 as long as the tube; staminal column pubescent at base; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Known only from the type (Rock 8595, A, BISH, CAS, GH, NSW, NY, S, W), collected in 1910 in subalpine shrubland at 2,130 m on Pu'unianiau, a small cinder cone on the northwestern slopes of Haleakalā, East Maui.

Although Rock (1913a, 1919a) treated this enigmatic plant as a species of Clermontia, it has little in common with that genus. On the basis of floral and vegetative morphology, it is better accommodated within Cyanea (Lammers, 1988b). In transferring it to that genus, a new name was necessitated by the prior existence of Cyanea haleakalaensis. The new epithet is the Hawaiian word pohaku, rock. Pohaku was used as a nickname for J.F.Rock, monographer of the Hawaiian Lobelioideae (Rock, 1919a), and the only scientist to ever see the species alive.

37. Cyanea procera Hillebr.

[Delissea procera (Hillebr.) St. John]

(end, E*)

Palm-like trees 3-9 m tall. Leaves oblanceolate, blades 60-75 cm long, 10-17 cm wide upper surface glabrous, lower surface pubescent at least along the veins, margins minutely callose-denticulate, apex obtuse, sessile. Inflorescences 10-20-flowered, peduncles 25-40 mm long, pedicels 6-10 mm long; hypanthium obconical, 15-20 mm long, 8-13 mm wide; calyx lobes shallowly deltate to shallowly triangular, 3-4 mm long, 4-5 mm wide, apex acute; corolla purplish, 60-80 mm long, 6-11 mm wide, glabrous, the tube suberect to slightly curved, the lobes 1/4-1/2 as long as the tube, all curved downward in the same plane, the corolla thus appearing unilabiate; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries ellipsoid to obovoid, 30-45 mm long, 20-28 mm wide. Formerly occurring in wet forest, ca. 600 m, eastern Moloka'i. Last collected in 1938 (Cranwell 3514, BISH, GB).

38. Cyanea profuga C. Forbes

[Delissea profuga (C. Forbes) St. John]

(end, Ex)

Unbranched shrubs 1.8-2.4 m tall. Leaves elliptic, blades 18-24 cm long, 7-10 cm wide, glabrous, margins callose-dentate, apex acuminate, base cuneate, petioles 8-12.5 cm long. Inflorescences 9-12-flowered, peduncles 25-40 mm long, pedicels 8-10 mm long; hypanthium oblong to obconical, 6-7 mm long, 3-4 mm wide; calyx lobes oblong, 3-5 mm long, 1-2 mm wide, apex rounded and apiculate; corolla white, 30-35 mm long, 2-3 mm wide, glabrous, the tube suberect, the lobes spreading, 3/4 as long to as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Known only from the type

page 461

(Forbes 313.Mo, BISH, MO, NY, UC) and one other specimen (Forbes 240.Mo, BISH), collected in 1912 at Mapulehu and Pelekunu, respectively, eastern Moloka'i.

39. Cyanea pycnocarpa (Hillebr.) F. Wimmer

[C. arborea Hillebr. var. pycnocarpa Hillebr.; Delissea pycnocarpa (Hillebr.) St. John]

(end, Ex)

Palm-like trees, height unknown. Leaves narrowly elliptic, blades ca. 34 cm long, ca. 4.5 cm wide, upper surface green, glabrous, lower surface pale green, pubescent along the midrib and veins, margins undulate and callose-denticulate, apex acuminate, base cuneate, petioles 3-4 cm long. Flowers sessile on peduncles 60-70 mm long; calyx lobes triangular in fruit, 4-5 mm long; corolla and stamens unknown. Berries ovoid, ca. 12 mm wide, crowned by the persistent calyx lobes. Known only from the type (Hillebrand s.n., B, presumably destroyed) collected in the Kohala Mountains, Hawai'i.

40. Cyanea quercifolia (Hillebr.) F. Wimmer

[C. solanacea Hillebr. var. quercifolia Hillebr.; Delissea quercifolia (Hillebr.) St. John]

(end, Ex)

Trees ca. 5 m tall; stems muricate to aculeate at least when young. Leaves broadly obovate, blades 30-50 cm long, 9-16 cm wide, upper surface papillose, lower surface pubescent, margins lobed or cleft toward apex and at middle, parted or divided at base, the lobes and apex rounded or obtuse, petioles ca. 4 cm long. Flowers (fide Hillebrand) as in Cyanea solanacea, except hypanthium and corolla glabrous and calyx lobes ca. 4 mm long. Known only from the type (Hillebrand 99, K) collected at 915-1,220 m on the leeward slopes of Haleakalā, East Maui.

This poorly known species is apparently related to Cyanea solanacea of Moloka'i, but differs in its distinctive leaf outline. The type of C. quercifolia var. atropurpurea (St. John et al. 11401, BISH, L) is a specimen of C. stictophylla.

41. Cyanea recta (Wawra) Hillebr.

[Delissea recta Wawra; Cyanea larrisonii Rock; C. rockii F. Wimmer; C. salicina H. Lév.; Delissea larrisonii (Rock) St. John; D. rockii (F. Wimmer) St. John]

(end, Ex)

Unbranched shrubs 1-1.5 m tall. Leaves narrowly elliptic, blades 12-28 cm long, 1.2-5 cm wide, upper surface green, glabrous, lower surface whitish green to pale green, glabrous or pubescent, margins minutely callose-denticulate, apex acuminate, base attenuate, petioles 1.5-4 cm long. Inflorescences 5-7-flowered, peduncles 7-10 mm long, pedicels 5-17 mm long; hypanthium obovoid, 6-10 mm long, 3-4 mm wide, pubescent; calyx lobes triangular, 2-2.5 mm long, 1-1.5 mm wide, apex acute; corolla purple or white with purple longitudinal stripes, 30-40 mm long, 3-4 mm wide, densely pubescent, the tube suberect, the lobes spreading; staminal column glabrous or sparsely pubescent at base; anthers minutely papillose, the lower 2 with apical tufts of white hairs. Berries purple, obovoid. Formerly occurring in mesic forest, 550-700 m, northeastern Kaua'i. Last collected in 1927 (MacDaniels 684, BISH).

42. Cyanea scabra Hillebr.

[C. holophylla Hillebr.; Delissea holophylla (Hillebr.) St. John; D. scabra (Hillebr.) St. John]

(end)

Shrubs 0.5-1.5 m tall; stems branched, smooth to muricate or aculeate. Leaves broadly elliptic to obovate, blades 15-23 cm long, 7-11 cm wide, upper surface green, glabrous to sparsely muricate, lower surface pale green, glabrous, pubescent muricate, or aculeate, margins callose-dentate, apex acuminate, base obtuse, petioles 4-7 cm long, glabrous, muricate, or aculeate. Inflorescences 6-12-flowered, peduncles 15-40 mm long, pedicels 12-18 mm long; hypanthium obconical, 4-6 mm long, 3-4 mm wide; calyx lobes oblong, 4-10 mm long, 2-3 mm wide, apex obtuse, truncate, or emarginate, sometimes with an apiculation; corolla white, often with pale lilac longitudinal stripes, 45-53 mm long, 4-7 mm wide, glabrous or pubescent, the tube arcuate, the lobes spreading, 1/4-1/3 as long as the tube, often muricate; staminal

page 462

column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellowish orange, ellipsoid, 10-13 mm long, crowned by the persistent calyx lobes. Occurring in mesic valleys, 460-850 m, West Maui. — Plate 55.

Because Cyanea scabra and C. holophylla are wholly sympatric and differ only in their muricate vs. smooth corolla lobes, they are regarded as conspecific. The varieties assigned to C. scabra by previous authors are interpreted here as synonyms of other taxa. Cyanea scabra var. sinuata is tentatively included in C. solanacea, while var. longissima is recognized at specific rank as C. longissima. The situation is more complex in var. variabilis. The holotype [Rock 10259 (coll. Lyon), BISH] is referred to Cyanea longissima, while the paratypes (Rock 8789a, 8797a, 8798, BISH) are referred to C. glabra.

43. Cyanea shipmanii Rock

[C. grimesiana Gaud. var. citrullifolia A. Gray; Delissea shipmanii (Rock) St. John]

(end, E*)

Shrubs 2.5-4 m tall; stems unbranched or sparingly branched at base, muricate, more so in juveniles. Leaves pinnately divided, cut 3/4-7/8 the distance to the midrib, blades 17-30 cm long, 7-14 cm wide (across the sements), upper surface glabrous, lower surface pubescent on the midrib and veins, muricate in juveniles, segments 10-15 per side, oblanceolate, pinnately divided, petioles 3-5.5 cm long, muricate. Inflorescences 10-15-flowered, pubescent, peduncles 10-30 mm long, pedicels 10-15 mm long; hypanthium obconical, 4-6 mm long, 3-4 mm wide, glabrous; calyx lobes oblong, 3-6 mm long, 0.5-1 mm wide, apex acute; corolla pale greenish white, 30-36 mm long, 3-4 mm wide, the tube gently curved, the lobes spreading, approximately as long as the tube; staminal column densely pubescent; anthers densely pubescent, the lower 2 with apical tufts of white hairs. Berries broadly ellipsoid, 12-15 mm long, 11-13 mm wide. Occurring in mesic forest, 1,800-1,900 m, windward slopes of Mauna Kea, Hawai'i.

Cyanea grimesiana var. citrullifolia is based upon a sterile specimen (U.S. Expl. Exped. s.n., GH, US) collected in 1840 on Mauna Kea, Hawai'i. The leaves are aculeate and pinnately divided into 10-15 pairs of pinnately divided segments. Rock (1919a) regarded these as juveniles of C. tritomantha. However, true juveniles of that species (e.g., Carlquist 2245b, BISH, L, MICH, RSA; Lammers et al., 5802, OS) bear muricate undivided leaves with dentate to shallowly lobed margins. Cyanea grimesiana var. citrullifolia thus is best regarded as a juvenile of C. shipmanii.

44. Cyanea solanacea Hillebr.

[C. ferox Hillebr.; C. f. var. laevicalyx Skottsb.; C. scabra Hillebr. var. sinuata Rock; C. s. var. variabilis Rock f. sinuata (Rock) F. Wimmer; Delissea ferox (Hillebr.) St. John; D. f. var. laevicalyx (Skottsb.) St. John; D. solanacea (Hillebr.) St. John]

(end) Pōpolo, hāhā nui

Shrubs 2-5 m tall; stems branched, muricate to aculeate (densely so in juveniles). Leaves dimorphic: juveniles pinnately cleft to divided, blades 20-40 cm long, 6-13 cm wide (across the segments), upper surface green, papillose to aculeate, lower surface pale green, pubescent and muricate to aculeate, segments 9-13 per side, oblong to oblanceolate or triangular, 10-25 mm wide, margins of the segments irregularly lobed to divided, apex acute or obtuse, petioles 2.5-5 cm long, pubescent and muricate to aculeate; adults elliptic to oblanceolate or obovate, blades 16-36 cm long, 4-14 cm wide, upper surface green, glabrous or papillose, lower surface pale green, pubescent, margins irregularly callose-dentate, lobed, or cleft, apex acuminate or acute, base cuneate, petioles 2.5-6 cm long, glabrous or pubescent. Inflorescences 5-10-flowered, peduncles 20-50 mm long, pedicels 8-12 mm long; hypanthium oblong to obconical, 8-10 mm long, 3-4 mm wide, glabrous, pubescent, or muricate; calyx lobes oblong or ovate, 4-15 mm long, 3-6 mm wide, apex obtuse to truncate or emarginate, sometimes with an apiculation; corolla white tinged with pale lilac, 48-60 mm long, 4-5 mm wide, pubescent or rarely subglabrous, the tube suberect to gently curved, the lobes spreading, 3/4 as long to as long as the tube; staminal column pubescent; anthers pubescent, the lower 2 with apical tufts of white hairs. Berries orange, ellipsoid, 11-16 mm long, crowned by the

page 463

page 464

persistent calyx lobes. Occurring in wet forest, 1,080-1,190 m, Moloka'i and possibly West Maui.

Dissection of the leaf margins varies not only with age, but also with geography. Plants of northern Moloka'i (Wai'ale'ia to Kamakou) tend to have leaves more deeply cut than do plants of southern Moloka'i (Kamolo to Pauwalu). The former have been called Cyanea ferox, the latter, C. solanacea. Because of this overlap between geographic and developmental variation, recognition of infraspecific taxa on this basis is inadvisable. Cyanea scabra var. sinuata of West Maui, known only from the type (Hillebrand 5, B, now destroyed), is tentatively included in C. solanacea. It is not at all similar to C. scabra and its allies, but rather resembles narrow-leaved specimens of C. solanacea from Oloku'i Plateau, Moloka'i (e.g., Higashino 9445 & 9464, BISH). A few sterile specimens from Pu'ukukui, West Maui (e.g., Hobdy 660, BISH; Warshauer 3070, BISH), may also belong here.

45. Cyanea solenocalyx Hillebr.

[C. ovatisepala F. Wimmer; C. solenocalyx f. glabrata (Rock) Degener; C. s. var. glabrata Rock; C. s. var. latifolia F. Wimmer; C. wailauensis Rock; Delissea ovatisepala (F. Wimmer) St. John; D. solenocalyx (Hillebr.) St. John; D. s. var latifolia (F. Wimmer) St. John; D. wailauensis (Rock) St. John]

(end) Pua kala

Palm-like trees 1-4 m tall; stems unarmed or muricate, aculeate in juveniles. Leaves dimorphic: juveniles ovate to broadly elliptic, blades 8-30 cm long, 6-18 cm wide (across the lobes), both surfaces pubescent and muricate to aculeate, margins irregularly lobed or cleft, apex acute to acuminate, base rounded to truncate or subcordate, petioles 3.5-11 cm long, muricate to aculeate; adults oblong to obovate, blades 25-60 cm long, 9-20 cm wide, upper surface glabrous, lower surface pubescent, unarmed or muricate along the midrib, margins callose-denticulate, sometimes shallowly lobed, apex acute to obtuse, base cuneate to rounded, petioles 8.5-16 cm long, muricate. Inflorescences 5-10-flowered, peduncles 30-50 mm long, pedicels 12-24 mm long; hypanthium obconical, 10-15 mm long, 5-10 mm wide at apex; calyx lobes oblong to narrowly oblong, 15-30 mm long, 5-8 mm wide and connate 5-27 mm from the base, or 3-4 mm wide and distinct, or ovate and 5-8 mm long, 4-5 mm wide; corolla white with a median purple longitudinal stripe on each lobe, or heavily suffused with purple, 34-45 mm long, 5-10 mm wide, glabrous or pubescent, the tube suberect, the lobes erect, ca. 1/4 as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellow to yellowish orange, ovoid, 16-18 mm long. Occurring in wet forest, 900-1,390 m, eastern Moloka'i. — Plate 55.

46. Cyanea spathulata (Hillebr.) A. Heller

[C. coriacea (A. Gray) Hillebr. var. spathulata Hillebr.; C. spathulata subsp. longipetiolata Lammers; Delissea spathulata (Hillebr.) St. John; Rollandia fauriei H. Lév.]

(end)

Branched shrubs 2-3 m tall. Leaves narrowly oblanceolate to narrowly elliptic, blades 8-22 cm long, 1-3 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or sparsely pubescent, margins minutely callose-crenulate, often revolute, apex obtuse to acute, base attenuate, narrowing into a winged petiole 1.5-3.5(-6) cm long, or narrowly cuneate with non-winged petioles 5.5-12 cm long. Inflorescences 8-10-flowered, the rachis condensed or obsolete, peduncles 15-50 mm long, pedicels 8-15 mm long; hypanthium broadly ellipsoid to broadly obovoid, 2-3 mm long, 1.5-2.5 mm wide; calyx lobes dentiform, 0.5-1 mm long, apex acute to acuminte; corolla white or greenish white, sometimes tinged pale purplish, 15-30 mm long, 1.5-3 mm wide, glabrous, the tube suberect, the lobes spreading; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries purple, globose, 5-8 mm wide. Occurring in diverse mesic forest to wet forest, 700-1,200 m, western and south-central Kaua'i.

Two subspeces may be distinguished: subsp. longipetiolata with the corolla 20-30 mm long, leaves narrowly elliptic, the bases narrowly cuneate, and petioles non-winged, 5.5-12 cm long (Mount Kāhili, 700-800m,

page 465

south-central Kaua'i); and subsp. spathulata [incl. Rollandia fauriei] with the corolla 15-23 mm long, leaves narrowly oblanceolate, the bases narrowly attenuate into a winged petiole 1.5-3.5(-6) cm long ('Ōpaewela and Kahōluamanu to Kōke'e and Kalalau, 900-1,200 m, western Kaua'i).

47. Cyanea stictophylla Rock

[C. nelsonii St. John; C. palakea C. Forbes; C. quercifolia (Hillebr.) F. Wimmer var. atropurpurea F. Wimmer; C. stictophylla var. inermis Rock; Delissea nelsonii (St. John) St. John; D. palakea (C. Forbes) St. John; D. quercifolia (Hillebr.) St. John var. atropurpurea (F. Wimmer) St. John; D. stictophylla (Rock) St. John; D. s. var inermis (Rock) St. John]

(end, E*)

Shrubs or trees 0.6-6 m tall; stems sparingly branched, unarmed or muricate. Leaves narrowly oblong to oblanceolate, blades 20-38 cm long, 4-8 cm wide, glabrous or muricate, margins callose-denticulate and shallowly lobed, apex acute, base cuneate, petioles 1.5-3.5 cm long, pubescent. Inflorescences 5-6-flowered, pubescent, peduncles 10-40 mm long, pedicels 7-22 mm long; hypanthium ovoid to obconical, 5-6 mm long, sparsely pubescent; calyx lobes triangular, 2-4 mm long, 1-2 mm wide, apex acuminate; corolla yellowish white or purple, 35-50 mm long, 5-6 mm wide, sparsely pubescent, the tube arcuate, the lobes spreading, 1/2-3/4 as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries orange, globose. Occurring in mesic to wet forest, 1,400-1,950 m, Kona coast and Ka'ū District, Hawai'i. This species has been collected only twice since 1958: in 1972 (Jacobi 105, BISH) and in 1982 (Warshauer et al. 3602, BISH).

48. Cyanea superba (Cham.) A. Gray

[Lobelia superba Cham.; Cyanea regina (Wawra) Rock; C. superba subsp. regina (Wawra) Lammers; C. s. var. regina (Wawra) Hillebr.; C. s. var. velutina Rock; Delissea regina Wawra; D. superba (Cham.) St. John; D. s. var. velutina (Rock) St. John; Macrochilus superbus (Cham.) K. Presl]

(end, E*)

Palm-like trees 4-6 m tall. Leaves oblanceolate, blades 50-100 cm long, 10-20 cm wide, glabrous or lower surface densely pubescent, margins callose-crenulate, apex acute, base cuneate, petioles 5-8 cm long. Inflorescences 5-15-flowered at apex, pendent, hanging below the leaves, densely pubescent, peduncles 200-350 mm long, pedicels 2-6 mm long, bracts oblanceolate, 12-45 mm long, apex obtuse to acute; hypanthium obconical, 10-16 mm long, densely pubescent; calyx lobes narrowly oblong, 6-16 mm long; corolla whitish or cream with purple veins, 55-80 mm long, 7-11 mm wide, densely pubescent, the tube curved, the lobes 3/4 as long to as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries yellow to orange, obovoid, 16-22 mm long, 10-16 mm wide. Occurring in mesic forest, 580-610 m, O'ahu. — Plate 55.

Two subspecies may be distinguished: subsp. regina with peduncles 200-240 mm long, calyx lobes 6-10 mm long, corolla cream with purple veins, and floral bracts 12-36 mm long (southern Ko'olau Mountains); and subsp. superba [incl Cyanea superba var. velutina] with peduncles 300-350 mm long, calyx lobes 10-16 mm long, corolla whitish to cream, and floral bracts 24-45 mm long (northern Wai'anae Mountains). The persistence of the Wai'anae race has been documented several times in recent years (e.g., Herbst 2243, F, PTBG; Obata 76301, BISH), but the Ko'olau race has not been collected since 1932 (Russell 188, BISH). The chromosome number reported by de Vilmorin and Simonet (1972) (as Lobelia superba) of 2n = 42 probably is an error, and perhaps based upon another plant.

49. Cyanea sylvestris A. Heller

[Clermontia pluriflora St. John; Cyanea eriantha Skottsb.; C. feddei H. Lév.; C. knudsenii Rock; C. sylvestris var. eriantha (Skottsb.) F. Wimmer; Delissea knudsenii (Rock) St. John; D. sylvestris (A. Heller) St. John; D. s. var. eriantha (Skottsb.) St. John]

(end)

Branched or unbranched shrubs 1.5-6 m tall. Leaves obovate to oblanceolate, blades 18-37 cm long, 5-13 cm wide, upper surface green, glabrous, lower surface pale green, glabrous or pubescent, margins minutely callose-denticulate, apex obtuse,

page 466

acute or acuminate, base cuneate, petioles 2.5-8 cm long. Inflorescences 6-12-flowered, peduncles 10-40 mm long, pedicels 5-15 mm long; hypanthium obovoid to obconical, 6-10 mm long, 4-8 mm wide, pubescent; calyx lobes narrowly triangular, 2-5 mm long, 1-2 mm wide, apex acuminate or acute; corolla white with purple longitudinal stripes, 40-48 mm long, 4-5 mm wide, densely pubescent, the tube gently curved, the lobes spreading; staminal column glabrous or pubescent at base; anthers minutely papillose, sometimes sparsely pubescent, the lower 2 with apical tufts of white hairs. Berries yellow to orange, obovoid, 12-20 mm long, 7-10 mm wide. Occurring in diverse mesic forest, 200-1,050 m, Kaua'i.

50. Cyanea tritomantha A. Gray

[C. magnifica F. Wimmer; C. submuricata F. Wimmer; Delissea magnifica (F. Wimmer) St. John; D. submuricata (F. Wimmer) St. John; D. tritomantha (A. Gray) St. John]

(end, V) 'Akū

Palm-like trees 1.8-3 m tall. Leaves oblong to oblanceolate or obovate, blades 38-95 cm long, 11-20 cm wide, upper surface green, glabrous, lower surface pale green, pubescent and muricate along the midrib, margins callose-denticulate, apex acuminate to acute or obtuse, base cuneate to attenuate, petioles 2-18 cm long, muricate. Inflorescences 5-20-flowered, clustered among the leaves, peduncles 20-75 mm long, pedicels 12-30 mm long; hypanthium obconical, 10-14 mm long, 5-8 mm wide; calyx lobes narrowly triangular, 10-25 mm long, 1.5-3 mm wide, apex acuminate, apiculate; corolla blackish purple or greenish white externally, pale bluish white or white within, 60-75 mm long, 5-7 mm wide, densely pubescent, the tube suberect to slightly curved, the lobes spreading, 1/2-3/4 as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries orange, ellipsoid, 15-25 mm long, 10-22 mm in diameter, crowned by the persistent calyx lobes. Occurring in wet forest, 350-1,080 m, Waipi'o Valley, windward Mauna Kea, and windward Mauna Loa, Hawai'i.

Rock (1919a) regarded Cyanea grimesiana var. citrullifolia as the juvenile form of C. tritomantha. However, that taxon is treated here as a synonym of Cyanea shipmanii. True juveniles of C. tritomantha are virtually indistinguishable from those of C. platyphylla, even though the 2 species are not closely related. Cyanea platyphylla often flowers precociously, while C. tritomantha does not. Many of the sites where this species occurs have been damaged by feral pigs.

51. Cyanea truncata (Rock) Rock

[Rollandia truncata Rock; Cyanea juddii C. Forbes; C. truncata var. juddii (C. Forbes) St. John; Delissea truncata (Rock) St. John; D. t. var. juddii (C. Forbes) St. John]

(end, E*)

Shrubs 0.3-2 m tall; stems unbranched or sparingly branched from base, muricate. Leaves obovate, blades 22-60 cm long, 10-26 cm wide, upper surface green and glabrous, lower surface pale green, pubescent and sparsely muricate, margins callose-denticulate, apex acute to acuminate, base cuneate to truncate, petioles 8-25 cm long, muricate. Inflorescences 8-40-flowered, horizontal to pendent, pubescent, peduncle plus rachis 40-280 mm long, pedicels 10-28 mm long, bracts narrowly oblong, 9-20 mm long, apex obtuse; hypanthium obconical, 5-6 mm long; calyx lobes narrowly oblong, 4-10 mm long, 2-4 mm wide, pubescent, apex obtuse; corolla white, striped or suffused with magenta, 32-42 mm long, 5-9 mm wide, pubescent, the tube curved, the lobes erect, 1/4-1/2 as long as the tube; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries orange, globose, 8-10 mm long. Occurring in mesic forest, 240-400 m, windward slopes of the Ko'olau Mountains, O'ahu. Collected only twice since 1931: in 1959 (Pang s.n., B, K) and in 1980 (K. nagata et al. 2176, BISH, HLA). — Plate 55.

Cyanea remyi Rock is known only from the holotype (Rémy 302bis, P), collected during 1851-1855 without specific locality on Kaua'i or Ni'ihau. The specimen is too fragmentary to allow adequate

page 467

characterization, but it may be referrable to Cyanea truncata.

52. Cyanea undulata C. Forbes

[Delissea forbesii St. John, non D. undulata Gaud.; D. lydgatei St. John, nom. illeg.]

(end, E*)

Unbranched shrubs 1.8-3.6 m tall. Leaves narrowly elliptic, blades 28-36 cm long, 3-5 cm wide, upper surface glabrous, lower surface pubescent, margins undulate, apex acuminate, base attenuate, narrowing imperceptibly into a winged petiole 3-5 cm long. Inflorescences 5-6-flowered, pubescent, peduncles ca. 43 mm long, pedicels ca. 10 mm long; hypanthium oblong, pubescent; calyx lobes triangular, ca. 4 mm long, apex acute; corolla (not fully expanded) yellowish, slightly curved, pubescent, the lobes spreading; staminal column pubescent; anthers glabrous. Berries yellow, obovoid, ca. 17 mm long, ca. 11 mm wide. Known only from the type (Forbes 292.K, BISH, K, US), collected in 1909 near Wahiawa Bog, south-central Kaua'i.

The prior existence of the combination Delissea undulata necessitates a new name for Cyanea undulata when treated as a species of Delissea. For this reason, St. John (1987b) published Delissea forbesii as an avowed substitute for Cyanea undulata. Apparently, he forgot that he had done so, since 4 months later (St. John, 1987c), he published Delissea lydgatei, also as an avowed substitute for Cyanea undulata. This second name is superfluous and thus illegitimate.

4. DELISSEA Gaud.

Terrestrial branched or unbranched shrubs or trees, rarely palm-like or subherbaceous, unarmed. Leaves toothed or irregularly laciniate, petiolate. Flowers 5-20 in axillary racemes; hypanthium ovoid, ellipsoid, cylindrical, or rarely campanulate; calyx lobes subulate to triangular, shorter than the hypanthium; corolla zygomorphic, white or greenish white, rarely suffused or striped with purple or blue, suberect to strongly curved, dorsally cleft to about the middle, with a small knob at the terminus of the cleft, sometimes with 2 additional lateral knobs, glabrous or rarely pubescent; stamens connate; lower 2 anthers with apical tufts of white hairs; ovary inferior, 2-celled; stigma 2-lobed. Fruit a purple berry. Seeds white, dull, transversely rugose, 0.7-1.5 mm long. [Chamisso, 1833a; Degener & Degener, 1982; Endlicher, 1836; Gaudichaud-Beaupré, 1829a; Gray, 1861b; Léveillé, 1913b; Mabberley, 1975a; Meyen, 1843; Rock, 1957; St. John, 1959, 1969b, 1977b, 1978d, 1980e, 1985a, 1985b, 1986b, 1987b, 1987c; St. John & Takeuchi, 1987; Skottsberg, 1926; Wawra, 1873b, 1873c]

An endemic genus of 9 species, arranged in 2 sections: Delissea [incl. sect. Micranthae (Hillebr. Rock] with suberect corollas 14-26 mm long; and Macranthae (Hillebr.) Rock with curved corollas 37-60 mm long. Some species may undergo dramatic changes in vegetative morphology during development, a phenomenonn well known in Cyanea (Carlquist, 1962a, 1969a). However, the paucity of extant taxa makes this difficult to verify. Several of the species are known only from Hillebrand's specimens and are presumed extinct. This lack of recent collections makes interpreting relationships within the genus and preparing detailed descriptions difficult. Although St. John (1987b) has greatly enlarged Delissea by merging it with Cyanea, the traditional circumscription of both genera is maintained here (see note under Cyanea). The genus commemorates M. Delisse, a physician from Mauritius, who served as naturalist of the D'Entrecasteaux expedition to the South Pacific in 1800-1804.

page 468

KEY TO THE SPECIES OF DELISSEA

1. Delissea fallax Hillebr.
(end, Ex)

Stems unbranched, leafy in the upper portion. Leaves oblong to oblanceolate, blades 14-24 cm long, 3-6 cm wide, margins denticulate, apex mucronate, base atttenuate, petioles 5-13 cm long. Peduncles 35-50 mm long, pedicels 8-10 mm long; hypanthium ovoid, 4-7 mm long; calyx lobes subulate, 1-3 mm long; corolla suberect, 15-18 mm long, with 1 dorsal knob; anthers glabrous. Berries subglobose, 6-8 mm in diameter. Known only from a few specimens collected by Hillebrand (Hillebrand s.n. GH; Hillebrand 82 & 82b, K) in Hāmākua and Hilo districts, Hawai'i.

2. Delissea laciniata Hillebr.
(end, Ex)

Stems branched. Leaves elliptic, blades 11-15 cm long, 3.5-5 cm wide (excluding lobes), irregularly laciniate, apex acute or acuminate, base attenuate or cuneate, the lobes triangular, 25-35 mm long, margins of the lobes irregularly dentate to serrulate, petioles 7-12 cm long. Inflorescences 6-10-flowered, peduncles 30-50 mm long, pedicels 10-12 mm long; hypanthium obovoid to turbinate; calyx lobes triangular to subulate, 1-3 mm long; corolla white, curved, 35-45 mm long, with 1 dorsal and 2 lateral knobs; anthers sparsely pubescent at base. Berries bluish, obovoid, ca 12 mm long. Known only from the type (Hillebrand s.n., BISH, GH, MEL, RSA), collected at Wailupe, O'ahu.

Two pre-Hillebrand specimens may also be referable to this species: Barclay 1311 (BM) and Kastalsky s.n. (LE). The former is labeled "tree 10 ft high, soil rich mould, flowers white, found in hilly situations, Woahu [O'ahu] July 1837." It differs from the type in its more deeply dissected leaves. The latter specimen is labeled simply "Ins. Sandwich."

page 469

3. Delissea lauliiana Lammers
[D. laciniata Hillebr. var. parvifolia Rock]
(end, Ex)

Leaves ovate, blades 6-7 cm long, 2-3 cm wide, margins serrate, apex acuminate, base cuneate, petioles 1.5-2 cm long. Corolla ca. 25 mm long, with 1 dorsal knob. Berries 6-7 mm long. Known only from the type (Hillebrand s.n., B), collected at Wailupe, O'ahu.

The type could not be located at B and is presumed to have been destroyed in 1943 (Hiepko, 1987). A second specimen (Mann & Brigham s.n., G), lacking precise locality data, may also be referable to this species. Although this taxon is imperfectly known, it clearly cannot be equated with Delissea laciniata. Aside from major differences in foliar features, the small corolla indicates that it belongs to sect. Delissea, while D. laciniata is a member of sect. Macranthae. Because it cannot be identified with any known species of sect. Delissea, it is reluctantly regarded as a distinct species. To avoid confusion with D. parviflora, the varietal epithet parvifolia, small-leaved, was not transferred to specific rank. The new epithet is derived from the Hawaiian work lau li'i, also meaning small-leaved.

4. Delissea parviflora Hillebr.
(end, Ex)

Stems branched, leafy in the upper portion. Leaves lanceolate to elliptic, blades 20-23 cm long, 2-3 cm wide (excluding lobes), irregularly laciniate, apex acute, the lobes triangular, 6-12 mm long, margins of the lobes irregularly dentate to serrulate, petioles 2.5-3.7 cm long. Peduncles 18-24 mm long, pedicels 6-7 mm long; calyx lobes triangular, ca. 1 mm long; corolla suberect, ca. 18 mm long, with 1 dorsal knob; anthers glabrous. Berries subglobose, ca. 8 mm long. Known only from specimens collected by Hillebrand (Hillebrand s.n., GH; Hillebrand 84, K) in the Kohala Mountains, Hāmākua District, and at 1,500 m on Mauna Kea, Hawai'i.

Delissea parviflora differs from D. fallax primarily in its laciniate, rather than denticulate, leave margins. The 2 species are apparently sympatric and their flowers are virtually identical. This suggests that they may represent juvenile and adult stages, respectively, of a single biological species. In the absence of confirming observations, the status quo is maintained.

5. Delissea rhytidosperma H. Mann
[D. kealiae Wawra]
(end, E*)

Branched shrubs 0.5-2.5 m tall; stems leafy in the upper portion. Leaves lanceolate to oblanceolate or elliptic, blades 8-19 cm long, 2-5.5 cm wide, margins sinuate, dentate, or serrulate, apex acute to acuminate, base attenuate, petioles 2-11 cm long. Inflorescences 5-12-flowered, peduncles 10-20 mm long, pedicels 8-13 mm long; hypanthium obovoid; calyx lobes subulate, 0.5-1 mm long; corolla greenish white, sometimes suffused with pale violet, suberect, 14-20 mm long, glabrous externally, minutely pubescent within, with 1 dorsal knob; staminal column glabrous; anthers pubescent at base. Berries dark purple, globose, 7-12 mm long. Seeds 0.7-1 mm long. [2n=28*.] Rare in diverse mesic forest, 300-1,000 m, Kauai'i. — Plate 56.

6. Delissea rivularis (Rock) F. Wimmer
[Cyanea rivularis Rock]
(end, Ex)

Stems 4-5 m long, unbranched or branched near the base, pubescent, with an apical rosette of leaves. Leaves elliptic to oblanceolate, blades 20-30 cm long, 3-8 cm wide, both surfaces pubescent, margins crenulate to attenuate, petioles 4-8 cm long, pubescent. Inflorescences 6-12-flowered, pubescent, peduncles 40-80 mm long, pedicels 10-15 mm long; hypanthium campanulate; calyx lobes triangular to subulate, 3-4 mm long; corolla white with blue longitudinal stripes, curved, 30-40 mm long, pubescent, with 1 dorsal knob; staminal column glabrous; anthers pubescent at base. Berries dark purple, globose, 10-15 mm in diameter. Seeds 0.7-1 mm long. Formerly occurring in diverse mesic forest, 1,300-1,400 m, Kaua'i. Last collected in 1916 (Hitchcock 15333, US).

Although placed in Cyanea by Rock (1913a, 1919a) and St. John (1973a), this species has the diagnostic corolla and seed features of Delissea and is best transferred to that genus (Wimmer, 1943).

page 470

page 471

7. Delissea sinuata Hillebr.
[D. s. subsp. lanaiensis (Rock) Lammers; D. s. var. lanaiensis Rock]
(end, Ex)

Stems subherbaceous, 0.6-1.2 m long, unbranched. Leaves elliptic, oblanceolate, or spatulate, blades 15-28 cm long, 4-7.5 cm wide, margins sinuate, denticulate, or dentate, apex acute, base attenuate, petioles 2.5-12 cm long. Peduncles 12-50 mm long, pedicels 9-13 mm long; hypanthium obovoid, 8-10 mm long; calyx lobes subulate, 2-3 mm long; corolla white, curved, 37-40 mm long, with 1 dorsal knob; anthers sparsely pubescent at base, Berries purple, ellipsoid, 7-10 mm long, 6-8 mm wide. Seeds ca. 1 mm long. Formerly occurring in diverse mesic forest, Wai'anae Mountains, O'ahu, and Lāna'i.

Two subspecies may be distinguished: subsp. lanaiensis with peduncles 12-15 mm long and elliptic or oblanceolate leaves 15-21 cm long on petioles 2.5-7 cm long (Lāna'i); and subsp. sinuata with peduncles 37-50 mm long and spatulate or oblanceolate leaves 25-28 cm long on petioles 7.5-12 cm long (O'ahu). The O'ahu race was last collected in 1937 (Fosberg 14214, POM), and the Lāna'i race in 1915 (Munro 19, BISH; Munro s.n., RSA, S). Both are presumed extinct.

8. Delissea subcordata Gaud.
[D. s. var. kauaiensis St. John D. s. var. obtusifolia Wawra; D. s. var. waialaeensis St. John; D. s. var. waikaneensis St. John; Lobelia subcordata (Gaud.) Endl.]
(end)

Branched or unbranched shrubs 1-3 m tall. Leaves ovate or ovate-lanceolate, blades 12-30 cm long, 6-17 cm wide, margins crenate, denticulate, serrate, or doubly serrate, the teeth incurved, or occasionally irregularly laciniate with 1-6 triangular lobes 10-18 mm long toward the base, apex obtuse, acute, or acuminate, base cordate, subcordate, or truncate, petioles 6-18 cm long. Inflorescences 6-18-flowered, glabrous, peduncles 40-100 mm long, pedicels 13-18 mm long; hypanthium ovoid to cylindrical; calyx lobes subulate, 0.5-1 mm long; corolla white or greenish white, curved, 45-60 mm long, with 1 dorsal knob; anthers glabrous. Berries ovoid, 12-16 mm long, 8-12 mm in diameter. Seeds ca. 1 mm long. Occurring in diverse mesic forest, 250-400 m, Ko'olau Mountains, and 450-550 m, Wai'anae Mountains, O'ahu. — Plate 56.

This species also may have occurred on Kaua'i (St. John, 1977b). However, the sole voucher for its occurrence there (Mann & Brigham s.n., BISH) appears to be mislabeled. The persistence of this plant in the Wai'anae Mountains is well documented (e.g., K. Nagata 1306, BISH, HLA; K. Nagata et al. 1589, HLA). However, it has not been collected in the Ko'olau Mountains since 1934 (Morley 135, BISH) and may be extinct there. Foliar characters such as size and outline of the leaf blades, size and shape of the marginal teeth, and the outlines of the apex and base are extremely variable. In the absence of correlated suites of characters with consistent geographic patterns, the recognition of infraspecific taxa seems unwarranted.

page 472

9. Delissea undulata Gaud.
[Cyanea argutidentata F. Wimmer; Delissea argutidentata (F. Wimmer) St. John; D. konaensis St. John; D. niihauensis St. John; D. undulata var. argutidentata (F. Wimmer) F. Wimmer; D. u. subsp. kauaiensis Lammers; D. u. subsp. niihauensis (St. John) Lammers; Lobelia undulata (Gaud.) Endl.]
(end, Ex)

Unbranched palm-like trees 2-10 m tall; stems woody, 1-5 cm in diameter, with a dense apical rosette of leaves. Leaves lanceolate, elliptic, ovate, or cordiform, blades 5-21 cm long, (3-)3.5-10 cm wide, margins undulate or flat, denticulate, crenate, or coarsely serrate, apex acute to acuminate, base cordate, truncate, or cuneate, petioles 2-15 cm long. Inflorescences 5-20-flowered, peduncles 5-50 mm long, pedicels 3-13 mm long; hypanthium ovoid to ellipsoid, 3-7 mm long; calyx lobes triangular to subulate, 0.8-2 mm long; corolla greenish white, suberect, 16-25 mm long, with 1 dorsal and 2 lateral knobs; staminal column and anthers glabrous. Berries ovoid to globose, 6-12 mm long. Seeds 1-1.5 mm long. Formerly occurring on Ni'ihau; in mesic forest near Hanapēpē Falls, Kaua'i; on dry pali and exposed cliffs, West Maui; and in mesic forest, western and southwestern slopes of Hualālai, 1,000 m, and Pu'ulehua on the southern flank of Mauna Loa, 1,650-1,750 m, Kona District, Hawai'i. — Plate 56.

Three subspecies may be distinguished: subsp. kauaiensis with ovate leaves, blades (7-)14-18 cm long, (3-)7-10 cm wide, margins flat and coarsely serrate, apex acuminate, base cuneate to truncate, and peduncles 5-20 mm long (Kaua'i); subsp. niihauensis with cordiform leaves, blades 5-11 cm long, 3.5-7 cm wide, margins flat and crenate, apex acute, base cordate, and peduncles 5-15 mm long (Ni'ihau); and subsp. undulata [incl. Cyanea argutidentata, Delissea konaensis] with elliptic to lanceolate leaves, blades 14-21 cm long, 4-8 cm wide, margins undulate and denticulate, apex acute, base cuneate, and peduncles 15-50 mm long (Maui, Hawai'i). The type of Delissea undulata var. serrulata (Wawra 1943, W) is a specimen of Cyanea elliptica. The Kaua'i race was last collected in 1895 (Heller 2430, A, AC, BM, E, F, G, GH, K, MIN, MSC, NY, P, UC, US). The Ni'ihau race was last collected ca. 1870 (Sinclair s.n., BISH). Maui populations of subsp. undulata were last reported at about the same time (Hillebrand, 1888; specimens subsequently destroyed). On Hawai'i, it is known to have persisted until at least as late as 1971 (Herbst 2141, BISH, PTBG).

5. HIPPOBROMA G Don.

[Isotoma (R. Br.) Lindl. sect. Solenanthis (Kunth) A. DC; Laurentia Adans. sect. Hippobroma (G. Don) Endl., nom. illeg.; Lobelia L. sect. Solenanthis Kunth]

Erect of decumbent annual or biennial herbs; stems unbranched or sparingly branched. Leaves lanceolate to oblanceolate, coarsely dentate to pinnately lobed, subsessile. Flowers solitary in the axils of the upper leaves; hypanthium turbinate, with conspicuous rim; calyx lobes linear, denticulate and ciliate; corolla white, salverform, the tube straight and entire, the lobes subequal and lanceolate; anthers connate, with an apical tuft of hairs; staminal column adnate to corolla tube from the base to just below the orifice. Fruit a loculicidal capsule dehiscent by 2 apical valves, pendent. Seeds numerous, pale brown, minute, ellipsoid, minutely pitted-reticulate. [St. John & Hosaka, 1932]

A monotypic genus originally endemic to the West Indies. Wimmer (1953) treated its sole species as a member ofo Laurentia sect. Isotoma (R. Br.) Endl. [Isotoma (R. Br.) Lindl.], an otherwise Australasian group. However, its resemblance to these plants is largely superficial and it is best regarded as a distinct genus (McVaugh, 1940; Melville, 1960). Name derived from the Greek hippos, horse, and bromos, poison, in reference to its extremely toxic latex.

1. Hippobroma longiflora (L.) G. Don
[Lobelia longiflora L.; Isotoma longiflora (L.) K. Presl; Laurentia longiflora (L.) Endl., non Schlechter]
(nat) Star-of-Bethlehem, pua hōkū

Stems coarse, fleshy, 15-50(-80) cm long, glabrous or pubescent. Leaves 6-25 cm long, 1.5-8 cm wide, glabrous or pubescent on the veins, apex obtuse to acute, mucronulate. Pedicels erect at anthesis but declined in fruit, 5-10 mm long, pubescent; hypanthium 10-nerved, 8-11 mm long, 3-5 mm wide, sparsely pubescent; calyx lobes 10-22 long; corolla 7-15 cm long, pubescent externally, glabrous within; anthers white. Capsules ellipsoid, 1.8-2.5 cm long, 1-1.5 cm wide, pendent. Seeds 0.6-0.8 mm long. [2n=28.] Originally endemic to the West Indies, now naturalized as a weed throughout much of the tropics; in Hawai'i naturalized in low elevation disturbed areas with moderate rainfall on Kaua'i, O'ahu, Maui, and Hawai'i. First collected on Kaua'i in 1864-1865 (Mann & Brigham s.n., BISH). — Plate 56.

page 473

6. LOBELIA L.

[Dortmanna J. Hill; Galeatella (F. Wimmer) Degener & I. Degener; Neowimmeria Degener & I. Degener]

Unbranched or sparingly branched shrubs (endemic species) or annual or perennial herbs (introduced species). Leaves entire or callose-toothed. Flowers (20-)50-200(-400) in terminal racemes; corolla zygomorphic, suberect to arcuate, dorsally cleft almost to base; anthers connate; ovary inferior, 2-celled; stigma 2-lobed. Fruit a loculicidal capsule, dehiscent at apex by 2 valves. Seeds numerous, smooth. [Croizat, 1941; Degener & Degener, 1965a, 1974c; Fosberg & Hosaka, 1938; Gray, 1861b; Heller, 1987; Kuntze, 1981a; Mabberley, 1974, 1975a, 1975b; Rock, 1917b, 1918a; St. John, 1931, 1939b, 1985a, 1987d, 1987e; St. John & Hosaka, 1935, 1938; Skottsberg, 1926, 1928, 1944a, 1953; Wimmer, 1948]

A cosmopolitan genus of over 350 species. Wimmer (1948, 1953) treated the 13 endemic Hawaiian species as 2 sections of subg. Tupa (G. Don) F. Wimmer: Galeatella F. Wimmer with corolla crimson or yellowish to greenish white, 35-90 mm long, 8-20 mm wide, tube curved to arcuate, lobes spreading, anthers with apical tufts of white hairs on all 5, and seeds 1-3 mm long; and Revolutella F. Wimmer with corollas blue or magenta, 15-45 mm long, 3-5 mm wide, tube suberect, lobes spirally recurved, anthers with apical tufts of white hairs on the lower 2, and seeds ca. 0.5 mm long. Mabberley (1974, 1975b) suggested that these 2 groups might better be included in sect. Rhynchopetalum (Fesen.) Benth.: Galeatella as a subsection, and Revolutella as a series within subsect. Nicotianifoliae Mabb. Recognition at generic rank as Galeatella and Neowimmeria (Degener & Degener, 1965, 1974c) would be inadvisable without a fuller understanding of relationships within the entire genus. The morphological distinctness of the 2 groups suggests independent colonization of Hawai'i (Rock, 1919a; Skottsberg, 1928; Mabberley, 1975b). Revolutella is probably derived from tropical Asian species such as Lobelia nicotianifolia Heyne ex Roem. & Schult., while Galeatella may be derived from Brazilian species such as L. organensis Gardn. (Mabberley, 1975b) or from eastern Asian species such as L. boninensis Koidz. (Koidzumi, 1914). The generic epithet honors Flemish botanists Mathias de L'Obel (1538-1616).

page 474

KEY TO THE SPECIES OF LOBELIA

1. Lobelia dunbariae Rock
[L. costata F. Wimmer; L. dunbariae subsp. paniculata (Rock) Lammers; L. hillebrandii Rock var. paniculata Rock; Neowimmeria costata (F. Wimmer) Degener & I. Degener; N. dunbariae (Rock) Degener & I. Degener; N. paniculata (Rock) Degener & I. Degener]
(end, R)

Stems woody, decumbent, 3-8 dm long, leafy in upper 1/2. Leaves linear or narrowly elliptic, 8-20 cm long, 0.5-2 cm wide, glabrous, slightly paler on lower surface, margins minutely callose-denticulate, flat, apex acute to acuminate, sessile to subsessile. Inflorescences 2-6-branched at base, 15-25 cm long, pedicels 8-30 mm long, glabrous, bracts linear and 12-15 mm long, or narrowly elliptic and 30-60 mm long; hypanthium obconical, 4-10 mm long; calyx lobes linear and 3-5 mm long, or narrowly triangular to narrowly oblong and 14-16 mm long; corolla blue, 18-26 mm long, 3-4 mm wide, the lobes spirally revolute, staminal column sparsely pubescent; anthers glabrous, the lower 2 with apical tufts of white hairs. Capsules ovoid, 6-12 mm long. Seeds brownish, 0.5 mm long. Occurring on cliffs in wet forest, 1,000-1,200 m, Moloka'i — Plate 57.

Two subspecies may be distinguished: subsp. dunbariae with floral bracts 30-60 mm long, calyx lobes 14-16 mm long, and pedicels 15-30 mm long (Waihānau Stream); and subsp. paniculata [incl. Lobelia costata] with floral bracts 12-15 mm long, calyx lobes 3-5 mm long, and pedicels 8-12 mm long (Kawela, Waialea, and Waikolu). Although the latter can still be found (e.g. Lammers et al. 5706, OS), the former is known only from the type [Rock 13125 (coll. Dunbar), BISH), collected in 1918.

page 475

page 476

2. Lobelia erinus L.
(nat) Edging lobelia

Stems herbaceous, 1-2.5 dm long. Leaves 0.5-2 cm long, lower ones ovate, obovate, or spatulate, upper one linear. Calyx lobes subulate, 2-8 mm long; corolla blue, rose, or white, often with a white or yellow center, 8-16 mm long; staminal column glabrous; anthers sparsely pubescent, the upper 2 with apical tufts of white hairs. Capsules obconical, 5-8 mm long. Seeds brown, 0.3-0.5 mm long. [2n=14,28,42.] Native to South Africa, widely cultivated as an ornameental, with many named cultivars available; in Hawai'i cultivated and perhaps sparingly naturalized in disturbed sites. The only specimen examined [Degener 8016 (coll. Brumaghim), BISH] was collected in Hawai'i Volcanoes National Park, Hawai'i, in 1931.

Lobelia tenuior R. Br. of western Australia also is cultivated occasionally in Hawai'i (Neal, 1965). It may be distinguished from L. erinus by its taller stature (2-6 dm), 3-lobed lower leaves, and all 5 anthers with apical tufts of white hairs.

3. Lobelia gaudichaudii A. DC
[Dortmanna gaudichaudii (A. DC) Kuntze; Galeatella gaudichaudii (A. DC) Degener & I. Degener; G. g. var. koolauensis (Hosaka & Fosb.) Degener & I. Degener; Lobelia gaudichaudii var. coccinea Rock; L. g. subsp. koolauensis (Hosaka & Fosb.) Lammers; L. g. var. koolauensis Hosaka & Fosb.]
(end)

Stems woody, 3-10 dm long, 1.5-4 cm in diameters, unbranched, with a dense apical rosette of leaves. Leaves oblanceolate to oblong, 8-19 cm long, 1.3-2.8 cm wide, both surfaces glabrous or midrib pubescent on lower surface, margins thickened or revolute, callose-crenulate toward apex, ciliate toward base, apex acute, sessile. Inflorescences unbranched or rarely 2-6-branched from base, 40-70 cm long, glabrous, pedicels 25-50 mm long, bracts lanceolate to ovate, 18-32 mm long, 5-10 mm wide, glabrous, apex acute, base truncate; hypanthium obconical or obovoid, 4-10 mm long, glabrous; calyx lobes triangular, lanceolate, or ovate, 10-15 mm long, 3-5 mm wide, glabrous, apex acute; corolla crimson, rarely greenish or yellowish white, 50-75 mm long, 8-15 mm wide, glabrous, the tube curved, the lobes spreading; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Capsules ovoid, 15-20 mm long, 10-15 mm wide, with an acuminate conical apex. Seeds brownish, ovoid, compressed, minutely winged, 1-1.2 mm long, 0.5 mm wide. Occurring in cloudswept wet forest, 800-950 m, on the summit of the Ko'olau Mountains, O'ahu.

Two subspecies may be distinguished: subsp. gaudichaudii [incl Lobelia gaudichaudii var. coccinea] with corolla crimson and inflorescences usually unbranched (central and southern Ko'olau Mountains); and subsp. koolauensis with corolla greenish or yellowish white and inflorescences 2-6-branched (northern Ko'olau Mountains).

4. Lobelia gloria-montis Rock
[Galeatella gloria-montis (Rock) Degener & I. Degener; G. g. var. bryanii (St. John & Hosaka) Degener & I. Degener; G. g. var. kukuiensis (St. John & Hosaka) Degener & I. Degener; G. g. var. longibracteata (Rock) Degener & I. Degener; G. g. var. molokaiensis (Degener) Degener & I. Degener; G. g. f. sanguinea (St. John & Hosaka) Degener & I. Degener; G. longibracteata (Rock) Degener & I. Degener; Lobelia gaudichaudii A. DC var. albiflora St. John; L. g. var. gloria-montis (Rock) St. John & Hosaka; L. g. var. g. f. bryanii St. John & Hosaka; L. g. var. g. f. kukuiensis St. John & Hosaka; L. g. var. g. f. sanguinea St. John & Hosaka; L. g. var. longibracteata Rock; L. gloria-montis var. longibracteata (Rock) Rock; L. g. var. molokaiensis Degener; L. longibracteata (Rock) F. Wimmer]
(end)

Stems woody, 10-50 dm long, 2-7 cm in diameters, unbranched, with a dense apical rosette of leaves. Leaves oblanceolate to oblong, 8-24 cm long, 1.8-4 cm wide, both surfaces glabrous or midrib pubescent on lower surface, margins thickened or revolute, callose-crenulate toward apex, ciliate towards base, apex acute, sessile. Inflorescences unbranched or rarely 3-7-branched from base, 40-70 cm long glabrous or rarely pubescent, pedicels 25-50 mm long, bracts lanceolate to ovate, 25-75 mm long, 5-20 mm wide, glabrous, apex acute, base truncate; hypanthium obconical to obovoid, 4-10 mm long, glabrous or rarely pubescent; calyx lobes triangular to lanceolate or ovate, 15-35 mm long, 3-9 mm wide, glabrous or rarely pubescent, apex acute; corolla yellowish or greenish white, often with reddish purple longitudinal stripes, or rarley crimson, 75-90 mm long, 8-20 mm wide, glabrous or rarely minutely pubescent, the tube curved, the lobes spreading; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Capsules ovoid, 15-20 mm ong, 10-15 mm wide, with an acuminate conical apex. Seeds brownish, ovoid, compressed, minutely winged, 1-1.2 mm long, 0.5 mm wide. Occurring in bogs and wet forest at margins of bogs, 1,350-1,750m, Maui. Sterile specimens collected on Pāpa'alā Pali, Moloka'i (Degener 7778, NY; Warshauer 2390, BISH; Anderson 513, BISH), also appear referable to this species. — Plate 57.

page 477

5. Lobelia grayana F. Wimmer
[Lobelia neriifolia A. Gray, non Moris; Dortmanna neriifolia Kuntze; Neowimmeria grayana (F. Wimmer) Degener & I. Degener]
(end)

Stems woody, erect, 10-20 dm long, with a dense apical rosette of leaves. Leaves linear,10-24 cm long, 0.5-1.5 cm wide, upper surface glabrous, lower surface densely white tomentose except on the midrib, margins entire to minutely callose-denticulate, revolute, apex acute, sessile or subsessile. Inflorescences unbranched, 10-40 cm long, pedicels 8-12 mm long, tomentose, bracts subulate, 35-45 mm long; hypanthium obconical, 4-6 mm long, white tomentose; calyx lobes subulate, 6-10 mm long; corolla blue, 28-32 mm long, 3-5 mm wide, glabrous or pubescent, the lobes spirally revolute; staminal column pubescent; anthers glabrous or pubescent, the lower 2 with apical tufts of white hairs. Capsules obovoid, 9-10 mm long, white tomentose. Seeds brownish, 0.5 mm long. [2n=28*.] Occurring on cliffs in subalpine shrubland, 1,700-2,300 m, Haleakalā, East Maui.

6. Lobelia hillebrandii Rock
[Neowimmeria hillebrandii (Rock) Degener & I. Degener]
(end)

Stems woody, ascending, 4-8 dm long, leafy in upper 1/2. Leaves linear to narrowly elliptic, 10-22 cm long, 0.8-1.4 cm wide, upper surface glabrous, lower surface white tomentose, margins callose-serrulate, flat, apex acute, sessile or subsessile. Inflorescences unbranched, 8-22 cm long, pedicels 10-14 mm long, bracts linear, 10-14 mm long; hypanthium obconical, 3-4 mm long; calyx lobes subulate, 4-6 mm long; corolla blue to lilac, 18-24 mm long, 3-4 mm wide, glabrous, the lobes spirally revolute; staminal column pubescent; anthers glabrous, the lower 2 with apical tufts of white hairs. Capsules void, 5-6 mm long. Seeds brownish, 0.5 mm long. Occurring in wet forest, 450-1,500 m, Maui.

Lobelia hillebrandii var. paniculata of Moloka'i and L. h. var. monostachya of O'ahu are treated here as Lobelia dunbarii subsp. paniculata and L. monostachya, respectively.

7. Lobelia hypoleuca Hillebr.
[L. h. var. heterocarpa F. Wimmer; L. h. f. macrophyta Rock; L. h. var. Rockii St. John & Hosaka; Neowimmeria heterocarpa (F. Wimmer) Degener & I. Degener; N. hypoleuca (Hillebr.) Degener & I. Degener; N. h. var. macrophyta (Rock) Degener & I. Degener; N. rockii (St. John & Hosaka) Degener & I. Degener]
(end) Kuhi'aikamo'owahie, liua, mo'owahie, 'ōpelu

Stems woody, erect, 15-30 dm long, leafy in upper 1/2. Leaves narrowly elliptic to elliptic, 30-65 cm long, 2-10 cm wide, upper surface glabrous, lower surface densely white tomentose except on the midrib, margins callose-denticulate, revolute or flat, apex acute to acuminate, sessile or subsessile. Inflorescences 3-7-branched at base, 15-40 cm long, pedicels 12-15 mm long, tomentose, bracts subulate, 10-18 mm long; hypanthium obconical, 4-6 mm long, white tomentose; calyx lobes subulate, 4-7 mm long; corolla blue, 28-32 mm long, 3-5 mm wide, pubescent, the lobes spirally revolute; staminal column pubescent; anthers glabrous or pubescent, the lower 2 with apical tufts of white hairs. Capsules cylindrical to obovoid, 8-18 mm long, white tomentose. Seeds brownish, ca. 0.5 mm long [2n=28*.] Occurring in mesic to wet forest, 600-1,500 m, on all of the main islands except Ni'ihau and Kaho'olawe.

This is the most widespread species of Hawaiian Campanulaceae, occurring on the 6 largest islands. As might be expected, it is extremely variable morphologically, especially in size and shape of the leaves. However, no geographically correlated patterns are discernible within this variation and the recognition of infraspecific taxa is thus unwarranted.

page 478

8. Lobelia kauaensis (A. Gray) A. Heller
[L. gaudichaudii A. DC var. kauaensis A. Gray; Galeatella kauaensis (A. Gray) Degener & I. Degener]
(end) Pu'e

Stems woody, 10-20 dm long, 3-6 cm in diameter, unbranched, with a dense apical rosette of leaves. Leaves narrowly lanceolate or narrowly oblong, 12-30 cm long, 1.5-3 cm wide, glabrous, margins thickened, revolute, entire to minutely callose-crenuate, apex acute to acuminate, base narrowed, sessile. Inflorescences 2-5-branched from base, (40-)80-150 cm long, glabrous, pedicels 15-30 mm long, bracts narrowly lanceolate, 15-30 mm long, 4-8 mm wide, glabrous or margins ciliate, apex acuminate, base truncate; hypanthium obconical, 5-8 mm long; calyx lobes triangular to lanceolate, 8-12 mm long, 3-5 mm wide, glabrous, apex acute; corolla yellowish to greenish white with purplish longitudinal stripes, 45-60 mm long, 8-12 mm wide, the tube curved, the lobes spreading; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Capsules pyriform, 18-21 mm long, 13-15 mm wide, with an acuminate conical apex. Seeds brownish, ovoid, compressed, minutely winged, 2-3 mm long. Occurring in bogs, 640-1,575 m, Wahiawa Bog and Wai'ale'ale, Kaua'i. — Plate 57.

Plants at Wai'ale'ale tend to be smaller and more compact than those at Wahiawa Bog; ths is possibly correlated with elevational differences. It may also be attributable to introgression from Lobelia villosa, which also grows on Wai'ale'ale. Several specimens from this locality, including the type of L. gaudichaudii var. kauaensis f. hirsuta St.John (Rock 12845a, BISH), appear to represent hybrids between these 2 taxa.

9. Lobelia monostachya (Rock) Lammers
[L. hillebrandii Rock var. monostachya Rock; Neowimmeria monostachya (Rock) Degener & I. Degener]
(end Ex)

Stems woody, prostrate, 1.5-2.5 dm long, leafy in upper 1/2. Leaves linear, 7-15 cm long, 0.4-0.7 cm wide, glabrous, margins entire to minutely callose-denticulate, flat or revolute, apex acuminate, sessile or subsessile. Inflorescences unbranched, 10-15 cm long, glabrous, pedicels 12-18 mm long, bracts filiform, 2-7 mm long; hypanthium obconical, 2-3 mm long; calyx lobes subulate to linear, 1-2.5 mm long; corolla magenta, 15-18 mm long 4-5 mm wide, glabrous, the lobes spirally revolute; staminal column glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Capsules unknown. Formerly occurring on dry, exposed slopes in mesic forest, southern Ko'olau Mountains, O'ahu.

This species is known only from the type collected in Niu (Hillebrand & Lydgate s.n., BISH) and 3 additional specimens collected in Mānoa Valley during the 1920s (Rock s.n., BISH; Topping 3228, NY; Degener & Topping 7964, A). The flowers on the type are mere buds, but those on the Rock and Topping specimens are in full anthesis. This permitted a better characterization of the plant and the realization that it was not at all related to Lobelia hillebrandii, but rather to L. niihauensis.

page 479

10. Lobelia niihauensis St. John
[L. n. var. forbesii St. John; L. n. var. meridiana St. John; L. tortuosa A. Heller, non (Benth.) Kuntze; L. t. f. glabrata Skottsb.; L. t. var. haupuensis St. John; L. t. var. intermedia St. John; Neowimmeria intermedia Degener & I. Degener; N. meridiana (St. John) Degener & I. Degener; N. niihauensis (St. John) Degener & I. Degener; N. tortuosa Degener & I. Degener; N. t. var. glabrata Degener & I. Degener]
(end, E*)

Stems woody, prostrate, 2-4 dm long branched at or below the middle, each branch with an apical rosette of leaves. Leaves narrowly elliptic or oblanceolate, 7-15 cm long, 0.7-1.8 cm wide, glabrous or pubescent, margins entire to callose-serrulate, flat to undulate or revolute, apex acute to acuminate, sessile or subsessile. Inflorescences unbranched, 12-15 cm long, glabrous or pubescent, pedicels 8-25 mm long, bracts filiform, linear, or subulate, 2-10 mm long; hypanthium obconical to hemispherical, 2-5 mm long, glabrous or pubescent; calyx lobes linear to triangular, 1-5 mm long, glabrous or pubescent; corolla magenta, 22-36 mm long, 3-5 mm wide, glabrous or pubescent, the lobes spirally revolute; staminal column glabrous or sparsely pubescent; anthers glabrous, the lower 2 with apical tufts of white hairs. Capsules ovoid, 6-8 mm long, glabrous or pubescent. Seeds brownish, 0.5 mm long. Occurring on exposed mesic to dry cliffs, 125-725 m, on Ni'ihau, Kaua'i, and the northern Wai'anae Mountains, O'ahu. — Plate 57.

In treating Lobelia niihauensis and L. tortuosa as conspecific, it has been necessary to retain the former name for the species. The latter name, though earlier, is illegitimate, being a later homonym of Lobelia tortuosa (Benth.) Kuntze. Although there are recent specimens from Kaua'i and O'ahu, on Ni'ihau it is known only from the type (Stokes s.n, BISH, NY), collected in 1912.

11. Lobelia oahuensis Rock
[Neowimmeria oahuensis (Rock) Degener & I. Degener]
(end, E*)

Stems woody, erect, 10-30 dm long, with a dense apical rosette of leaves. Leaves elliptic, 40-60 cm long, 4-6 cm wide, upper surface glabrous, lower surface densely grayish or greenish hirsute, margins callose-denticulate, revolute, apex acuminate, sessile or subsessile. Inflorescences 3-5-branched from base, 100-150 cm long, pubescent, pedicels 10-20 cm long, bracts linear, 20-40 mm long, 2-3 mm wide, apex acuminate; hypanthium obconical to obovoid, 4-6 mm long, pubescent; calyx lobes linear, acute, 12-18 mm long, 2-3 mm wide; corolla pale blue, 42-45 mm long, 4-5 mm wide, pubescent or glabrous, the lobes spirally revolute; staminal column pubescent; anthers pubescent or glabrous, the lower 2 with apical tufts of white hairs. Capsules obovoid, 10-17 mm long, 8-10 mm wide, pubescent. Seeds brownish, 0.5 mm long. Occurring in cloudswept wet forest, 850-920 m, on the windswpet summit of the Ko'olau Mountains, O'ahu. This species had not been collected since 1933 (Hosaka 1156, BISH) when it was rediscovered in 1984 (Obata 84-464, BISH).

12. Lobelia remyi Rock
[Neowimmeria remyi (Rock) Degener & I. Degener]
(end, Ex)

Stems unknown. Leaves elliptic, 12-16 cm long, 1.4-2.6 cm wide, upper surface glabrous, lower surface grayish tomentose, apex acuminate, base acute, abruptly contracted into a distinct petiole 5-8 mm long. Inflorescences unbranched, 12-15 cm long, pedicels 12-14 mm long; hypanthium hemispherical, ca. 5 mm long; calyx lobes triangular, apex acute to acuminate. Known only from the type specimen (Rémy 298, P), collected in the early 1850s on O'ahu.

13. Lobelia villosa (Rock) St. John & Hosaka
[L. kauaensis (A. Gray) A. Heller var. villosa Rock; Galeatella villosa (Rock) Degener & I. Degener]
(end)

Stems woody, 6-12 dm long, 3-6 cm in diameter, unbranched, with a dense apical rosette of leaves. Leaves lanceolate to oblong, 9-14 cm long, 2.5-4 cm wide, upper surface glabrous or sparsely pubescent, lower surface pubescent on the midrib, margins thickened, revolute, entire, pubescent, apex obtuse, base narrowed to subcordate or cordate, sessile. Inflorescences 3-5-branched from base or rarely unbranched, 40-60 cm long, densely pubescent, pedicels 25-30 mm long, bracts ovate, 20-30 mm long, 14-20 mm wide, pubescent, entire, apex obuse, base often auriculate; hypanthium broadly obconical, 5-8 mm long, 10-14 mm wide, pubescent; calyx lobes ovate to oblong, 10-15 mm long, 4-6 mm wide, pubescent, margins ciliate, apex obtuse; corolla yellowish to greenish white with purplish veins, 35-45 mm long, 8-12 mm wide, glabrous, the tube curved, the lobes spreading; staminal column glabrous; anthers glabrous, all 5 with apical tufts of white hairs. Capsules pyriform, with an acuminate conical apex, pubescent. Seeds brownish, ovoid, compressed, minutely winged, 1.5-2.5 mm long. Occurring in bogs, 1,200-1,580 m, Alaka'i Swamp and Wai'ale'ale, Kaua'i.

page 480

14. Lobelia yuccoides Hillebr.
[Dortmanna yuccoides (Hillebr.) Kuntze; Neowimmeria yuccoides (Hiller.) Degener & I. Degener]
(end) Pānaunau

Stems woody, erect, 15-20 dm long, with a dense apical rosette of leaves. Leaves linear, 24-35 cm long, 0.5-1.5 cm wide, upper surface glabrous, lower surface white tomentose, margins entire or minutely callose-denticulate, revolute, apex acute to acuminate, sessile or subsessile. Inflorescences unbranched, 60-100 cm long, pedicels 8-12 mm long pubescent, bracts subulate, 20-30 mm long; hypanthium obconical, 4-6 mm long, pubescent; calyx lobes subulate, 36 mm long; corolla blue to lilac, 36-40 mm long, 3-5 mm wide, pubescent, the lobes spirally revolute; staminal column pubescent; anthers pubescent at base, the lower 2 with apical tufts of white hairs. Capsules obovoid to cylindrical, 10-12 mm long, pubescent. Seeds brownish, 0.5 mm long. [2n=28*.] Occurring on dry ridges and canyons in diverse mesic forest, 750-1,200 m, Kaua'i and the Wai'anae Mountains, O'ahu.

7. ROLLANDIA Gaud.

[Delissea Gaud. sect. Rollandia (Gaud.) Baill.]

Terrestrial, unbranched shrubs, rarely subherbaceous, unarmed or occasionally muricate, more so in juveniles than adults, leafy at apex. Leaves entire or callose-crenulate, those of juveniles usually more deeply toothed or shallowly lobed. Flowers (3-)5-15(-20) in axillary, racemose inflorescences; hypanthium obovoid to cylindrical; calyx lobes shorter than or equalling the hypanthium, distinct or connate; corolla zygomorphic, pale to dark magenta, often with lighter or darker longitudinal stripes, rarely white, greenish, or purple, laterally compressd, strongly sigmoid, dorsally cleft to about the middle, glabrous or pubescent, rarely muricate, the lobes subequal and spreading, less than 1/4 as long as the tube; stamens connate, adnate to the corolla 1/3-1/2 its length; ovary inferior, 2-celled stigma 2-lobed. Fruit a yellow, orange or purple berry. Seeds pale brown to black, smooth, shiny. [Chamisso, 1833a; Forbes, 1912; Fosberg, 1936a; Gaudichaud-Beaupré, 1829a, 1841-?1852; Heller,1897; Léveilé, 1913b; Mabberley, 1975a; Meyen, 1843; Rock, 1917b, 1918a; St. John 1939b, 1940a, 1940b, 1979e, 1980e, 1985a, 1987f; St. John & Hosaka, 1935; Skottsberg, 1926; Wawra, 1873d]

An endemic genus of 8 species, all but one, Rollandia parvifolia of Kaua'i, restricted to O'ahu. The genus is closely related to Cyanea and differs primarily in that the staminal column is adnate to the corolla tube. The name commemorates M. Rolland, gunnery chief of the Freycinet expedition of 1817-1820, who rendered great service to ornithology.

As this book went to press, 2 new species of Rollandia were published (St. John, 1987f): R. alba St. John & Takeuchi and R. obatae St. John. Without examing the types, it not possible to ascertain the correct placement of these novelties on the basis of the inadequate "diagnoses." Regrettably, they are excluded from further consideration except to note that they most likely can be accommodated within previously recognized taxa.

page 481

KEY TO THE SPECIES OF ROLLANDIA

1. Rollandia angustifolia (Hillebr.) Rock

[R. longiflora Wawra var. angustifolia Hillebr.; R. angustifolia var. ochreata F. Wimmer]

(end)

Stems woody, 1-1.5 m long. Leaves whitish on lower surface, linear to narrowly elliptic, blades 16-36 cm long, 1.5-4 cm wide, glabrous, margins callose-crenulate to subentire, apex acute to acuminate, petioles 1.5-4.5 cm long. Inflorescences 3-6-flowered, peduncles 15-40 mm long, pedicles 10-22(-30) mm long; hypanthium obconical to turbinate, 6-12 mm long; calyx lobes connate, forming a truncate or irregularly toothed or lobed sheath, 2-8 mm long; corolla dark magenta, 5-9 cm long, glabrous; staminal column glabrous to sparsely pubescent; anthers glabrous or sparsely pubescent, the lower 2 with apical tufts of white hairs. Berries globose. Occuring in exposed cloudswept wet forest along crests, less often in mesic valleys, 450-670 m, Ko'olau Mountains, O'ahu. — Plate 58.

2. Rollandia crispa Gaud.

[Cyanea rollandia A. Gray, nom. illeg.; Lobelia crispa (Gaud.) Endl., non Graham; Rollandia crispa var. muricata Rock; R. grandifolia Hillebr.]

(end, E*)

Stems fleshy, 0.3-1.3 m long. Leaves broadly obovate, blades 30-75 cm long, 9-16 cm wide, upper surface glabrous, lower surface glabrous or pubescent, margins undulate, callose-crenulate to entire, apex acute to acuminate, petioles 0.8-4 cm long. Inflorescences 3-8-flowered, pubescent, peduncles 20-30 mm long, pedicels 10-20 mm long; hypanthium obconical, 8-12 mm long, pubescent; calyx lobes ovate to oblong, 6-12 mm long, 3-6 mm wide, pubescent, apex obtuse to truncate, base often cordate and overlapping; corolla pale magenta with darker longitudinal stripes, 4-6 cm long, pubescent; staminal column pale purplish, glabrous; anthers dark purple, glabrous to sparsely pubescent, the lower 2 with apical tufts of white hairs.

page 482

page 483

Berries globose, 10 mm long. Occurring in mesic to wet forest, 185-720 m, Ko'olau Mountains, O'ahu. Collected only twice since 1946; in 1970 (Herbst & Bishop 1537, HAW) and in 1985 (K. Nagata et al. 3279, HLA).

3. Rollandia humboldtiana Gaud.

[Delissea racemosa H. Mann; Rollandia humboldtiana f. albida St. John; R. pedunculosa Wawra; R. racemosa (H.Mann) Hillebr.]

(end)

Stems woody, 1-2 m long. Leaves obovate to broadly elliptic, blades 18-45 cm long, 7-16 cm wide, upper surface glabrous, lower surface pubescent, margins callose-crenulate, apex acute to acuminate, petioles 3-4 cm long. Inflorescences 5-12-flowered, pendent, pubescent, peduncles 80-250 mm long, pedicels 10-15 mm long; hypanthium oblong to obconical, 8-12 mm long, pubescent; calyx lobes oblong, 4-10 mm long, pubescent, apex truncate; corolla dark magenta or rarely white, 6-7.5 cm long, pubescent; staminal column glabrous; anthers pale magenta, glabrous, the lower 2 with apical tufts of white hairs. Berries pale orangish yellow, ellipsoid to obovoid, 10-14 mm long. Occurring in wet forest, 300-915 m, Ko'olau Mountains, O'ahu. — Plate 58.

4. Rollandia lanceolata Gaud.

[Clermontia aspera F. Wimmer, pro parte; Cyanea aspera A. Gray; C. coronata F. Wimmer; Delissea ambigua (Cham.) K. Presl; D. calycina (Cham.) K. Presl; D. coronata (F. Wimmer) St. John; D. delessertiana (Gaud.) A. Gray; D. lanceolata (Gaud.) A. Gray; Lobelia ambigua Cham.; L. calycina Cham.; L. lanceolata (Gaud.) Hook & Arnott; Rollandia ambigua (Cham.) G. Don; R. bidentata St. John; R. calycina (Cham.) G. Don; R. c. var. kaalae (Wawra) F. Wimmer; R. delessertiana Gaud.; R. humboldtiana Gaud. var. tomentella Wawra; R. kaalae Wawra; R. lanceolata subsp. calycina (Cham.) Lammers; R. l. var. glaberrima F. Wimmer; R. l. var. grandifolia A. DC; R. l. var. kipapaensis Hosaka ex Fosb.; R. l. f. rockii St. John & Hosaka; R. l. var. tomentella (Wawra) F. Wimmer; R. l. var. tomentosa (Hillebr.) Drake; R. l. var. viridiflora Rock; R. scabra Wawra; R. waianaeensis St. John]

Stems woody, 1-3 m long, muricate in juveniles, becoming merely scabrous or nearly smooth with age. Leaves elliptic or oblanceolate, blades 15-60 cm long, 5.5-14 cm wide, upper surface glabrous, muricate in juveniles, lower surface pubescent or rarely glabrous, margins callose-crenulate, coarsely callose-dentate in juveniles, apex acute to acuminate, petioles 1-9 cm long, pubescent or muricate. Inflorescences 4-16-flowered, peduncles 20-100 mm long, pedicels 5-30 mm long; hypanthium obovoid, obconical, or ellipsoid, 6-12 mm long, glabrous or pubescent; calyx lobes triangular, ovate, or oblong, 1-10 mm long, glabrous or pubescent, apex acute, obtuse, or truncate; corolla pale to dark magenta, rarely pale greenish, often with lighter or darker longitudinal stripes, 5-8 cm long, glabrous or pubescent; staminal column glabrous or pubescent; anthers pubescent, all 5 or only the lower 2 with apical tufts of white hairs. Berries obovoid to obpyriform, 16-20 mm long. [2n = 28*; subsp. calycina.] Occurring in mesic valleys and wet forest, 300-915 m, Ko'olau and Wai'anae mountains, O'ahu. — Plate 58.

Two subspecies may be recognized: subsp. calycina [incl. Clermontia aspera pro parte, Cyanea aspera, C. coronata, Rollandia bidentata, R. humboldtiana var. tomentella, R. kaalae, R. lanceolata var. tomentosa, R. scabra, R. waianaeensis] with calyx lobes oblong to ovate, 4-10 mm long, apex obtuse to truncate, corolla and hypanthium pubescent, lower leaf surface densely pubescent with branches and clustered hairs (common in the Wai'anae Mountains, scarce in the Ko'olau Mountains); and subsp. lanceolata [incl. Lobelia ambigua, Rollandia delessertiana, R. lanceolata var. glaberrima, R. l. var. grandifolia, R. l. var. rockii, R. l. var. viridiflora] with calyx lobes triangular, 1-3 mm long, apex acute to obtuse, corolla and hypanthium glabrous, and lower leaf surface pubescent with simple hairs or rarely glabrous (Ko'olau Mountains). Several specimens from the Ko'olau Mountains, including the type of Rollandia lanceolata var. kipapaensis (Hosaka 1027, BISH), represent transitional forms connecting the 2 subspecies.

Clermontia aspera is based on a mixed collection (Degener & Ordonez s.n., W), comprising a detached inflorescence of Clermontia oblongifolia subsp. oblongifolia

page 484

and leaves of Rollandia lanceolata subsp. calycina. Cyanea coronata is not related to C. procera as suggested by Wimmer (1943); the type (Forbes s.n., BISH) is a specimen of Rollandia lanceolata subsp. lanceolata. Lobelia ambigua was long considered a name of uncertain application (Rock, 1919a). Wimmer (1943) subsequently treated it as a synonym of Rollandia humboldtiana. However, the type (Chamisso s.n., LE) is a specimen of R. lanceolata subsp. lanceolata. Rollandia bidentata is based upon a precociously flowering juvenile individual (St. John 11038, BISH, US) of R. lanceolata subsp. calycina. The type of Rollandia degeneriana F. Wimmer (Degener et al. 11484, A, B, G, K, MASS, MO, NY, RSA, US, W) appears to represent a hybrid between R. lanceolata and R. longiflora.

5. Rollandia longiflora Wawra

[R.lanceolata Gaud. var. brevipes F. Wimmer; R. sessilifolia Degener]

(end)

Stems woody, 1-3 m long, muricate in juveniles, becoming smooth with age. Leaves elliptic or oblanceolate, blades 30-55 cm long, 6-12 cm wide, glabrous, muricate in juveniles, margins entire or callose-crenulate, irregularly cleft or lobed in juveniles, apex acute, sessile or on petioles 0.3-3 cm long. Inflorescences 5-10-flowered, peduncles 30-60 mm long, pedicels 5-15 mm long; hypanthium obconical, 6-10 mm long, glabrous; calyx lobes connate into an irregularly toothed sheath 2-4 mm long; corolla dark magenta, 6-9 cm long; staminal column dark magenta, glabrous; anthers glabrous, the lower 2 with apical tufts of white hairs. Berries obpyriform, 10-12 mm long. Occurring in mesic to wet forest, 275-760 m, Ko'olau and Wai'anae mountains, O'ahu.

6. Rollandia parvifolia C. Forbes

(end,Ex)

Stems 1 m long. Leaves narrowly elliptic, blades 18-22 cm long, 2.3-4 cm wide, glabrous, margins entire, apex acuminate, petioles ca. 2.8 cm long. Inflorescences 5-6-flowered, peduncles 30-40 mm long, pedicels ca. 10 mm long; hypanthium cylindrical, ca. 8 mm long; calyx lobes distinct, oblong, ca. 1. cm long, obtuse; corolla purple, ca. 6.5 cm long; staminal column pubescent; anthers pubescent. Berries unknown. Known only from the type specimen (Forbes 103.K, BISH) collected in 1909 at Wai'oli Valley, Kaua'i.

7. Rollandia purpurellifolia Rock

(end,Ex)

Stems fleshy, 0.2-0.5 m long. Leaves narrowly oblong to spatulate, blades 12-35 cm long, 3-6.5 cm wide, upper surface glabrous, lower surface purple, glabrous or sparsely pubescent on the veins, margins callose-crenulate toward apex, entire toward base, apex subacute, subsessile or on petioles 0.5-0.8 cm long. Inflorescences 3-6-flowered, peduncles 10-25 mm long, pedicels 8-16 mm long; hypanthium cylindrical to obconical, 4-5 mm long; calyx lobes connate, forming an irregularly 2-lipped or 5-lobed sheath 7-10 mm long; corolla dark magenta, 6-8 cm long; staminal column magenta, glabrous; anthers pale purple, glabrous, the lower 2 with apical tufts of white hairs. Berries dark purple, ellipsoid to obovoid, 15-18 mm long. Known only from a few specimens collected at ca. 600m in the vicinity of Kaluanui Valley, Ko'olau Mountains, O'ahu. Last collected in 1935 (Degener et al. 11476, B, G, MASS, MICH, MO, W).

8. Rollandia st. johnii Hosaka

[R. s. var. obtusisepala F. Wimmer]

(end)

Stems woody, 0.3-0.6 m long. Leaves lanceolate to oblanceolate, blades 6-13 cm long, 1.5-2 cm wide, upper surface glabrous, lower surface pubescent on the veins, margins callose-crenulate, strongly revolute, apex subacute to acuminate, petioles 0.6-1.5 cm long, pubescent to muricate. Inflorescences 5-20-flowered, peduncles 5-15 mm long, pedicels 5-10 mm long; hypanthium 4-7 mm long, sparsely pubescent; calyx lobes oblong to lanceolate, 3-10 mm long, 2-3 mm wide, apex obtuse, acute, or acuminate; corolla white, the lobes suffused with pale violet on inner surface, 3-6 cm long, glabrous; staminal column glabrous or sparsely pubescent; anthers dark purple, glabrous, the lower 2 with apical tufts of white hairs. Berries unknown. Endemic to windy cloudswept wet forest, summit of the central Ko'olau Mountains, O'ahu. — Plate 58.

page 485

8. TREMATOLOBELIA A. Zahlbr.

[Trematocarpus A. Zahlbr., non Kütz.]

Terrestrial or rarely epiphytic, unbranched shrubs (0.5-)1-4 m tall; stems woody, solitary, erect, monocarpic, with an apical rosette of leaves. Leaf margins subentire to callose-crenuate. Flowers 50-400 in terminal racemes composed of 5-20 horizontal branches, the flowers erect and secund, or rarely flowers 15-40 in unbranched vertical racemes, the flowers ascending and secund; hypanthium subglobose; calyx lobes shorter than to equalling the hypanthium, rarely 1/2 again as long; corolla zygomorphic, scarlet to rose or pink, rarely white, the tube gently to strongly curved, dorsally cleft to the base, laterally cleft to about the middle, forming 2 lateral lobes, the 3 ventral lobes forming a trifid lip; stamens connate, exserted from corolla; lower 2 anthers with apical tufts of white hairs; ovary inferior, 2-celled; stigma 2-lobed. Fruit a fleshy 10-ribbed capsule, the parenchymatous exocarp upon decomposition exposing the sclerenchymatous endocarp perforated by few to numerous irregular pores through which the seeds disperse. Seeds yellowish to brown, void, compressed, winged, ca. 1 mm long, ca. 0.5 mm wide. [Degener & Degener, 1968a, 1974c, 1983; Gray, 1861b; Hemsley, 1892, 1893; Kuntze, 1891; Mabberley, 1975a; St. John, 1971a, 1982a, 1985a; Schönland, 1894; Skottsberg, 1926, 1944; Stapf, 1893; Zahlbruckner, 1891]

An endemic genus of 4 species. Wimmer (1953) regarded Sclerotheca A. DC, endemic to Rarotonga, Rapa, Tahiti, and the Marquesas, as its closest relative. However, the species of Sclerotheca are branched shrubs with solitary axillary flowers and bear little similarity to Trematolobelia. The genus is more likely related to Lobelia, from which it differs only in the peculiar mode of fruit dehiscence and seed dispersal (Rock, 1919a; Carlquist, 1962b; St. John, 1982a). In fact, one cannot rule out the possibility that it is derived from the endemic Hawaiian sect. Galeatella or shares a common ancestor with that group (Mabberly, 1975a). Name derived from the Greek trematos, hole (in reference to the perforated capsule), and the genus Lobelia, to which the type species, Trematolobelia macrostachys, was originally assigned.

KEY TO THE SPECIES OF TREMATOLOBELIA

page 486

1. Trematolobelia grandifolia (Rock) Degener
[T. macrostachys (Hook. & Arnott) A. Zahlbr. var. grandifolia Rock; T. kohalaensis St. John; R. rockii St. John var. hawaiiensis St. John; T. wimmeri Degener & I. Degener]
(end)

Stems 1-4 m long, 1-4 cm in diameter, glabrous or pubescent. Leaves green on upper surface, lower surface pale green, narrowly oblong, broadly oblanceolate, or spatulate, blades 20-44 cm long, 2-7 cm wide, margins subentire or faintly collose-crenulate, apex acute to obtuse, sessile or tapering imperceptibly into a winged petiole 2-5 cm long. Inflorescences composed of 5-10 horizontal branches 30-60 cm long, pedicels 1-35 mm long, bracts elliptic to lanceolate, 15-50 mm long, 3-15 mm wide; hypanthium 5-8 mm wide; calyx lobes erect or ascending, narrowly triangular to lanceolate, 5-9 mm long; corolla white, white with faint rose streaks, rose, or pink, 5.5-8 cm long, the tube and trifid lip gently curved downward, the 2 lateral lobes recurved upward, the corolla thus appearing bilabiate; staminal column 4-6 cm long, sparsely pubescent; anthers pubescent at base. Capsules subglobose, 13-15 mm in diameter (fresh exocarp), 10-13 mm in diameter (dried endocarp). Occurring on wet windswept cliffs and in wet forest and low, boggy wet forest, in the Kohala Mountains (925-1275 m), in the saddle between Mauna Loa and Mauna Kea (800-1500m), and in and around Hawai'i Volcanoes National Park (625-1200m), Hawai'i. Collected only twice since 1958; in 1966 (Degener & Degener 34720, MO, NY) and in 1977 (Warshauer s.n., BISH).

2. Trematolobelia kauaiensis (Rock) Skottsb.
[T. macrostachys (Hook. & Arnott) A. Zahlbr. var. kauaiensis Rock]
(end) Kolli'i

Stems 1.5-3 m long, 1-1.5 cm in diameter. Leaves dark green on upper surface, lower surface pale green, linear-elliptic to elliptic, blades 12-22 cm long, 1-3 cm wide, glabrous, margins callose-crenulate, apex acuminate, base cuneate, petioles 1.3-2.5 cm long. Inflorescences composed of 5-10 horizontal branches 20-58 cm long, pedicels 15-20 mm long, bracts lanceolate, 9-25 mm long; hypanthium 5-8 mm wide; calyx lobes spreading, oblong, 4-10 mm long, apex obtuse; corolla scarlet, 5-5.5 cm long, the tube, trifid lip, and 2 lateral lobes all strongly curved downward in the same plane, the corolla thus appearing unilabiate; staminal column 4-5.5 cm long, sparsely pubescent; anthers pubescent in the lower 1/2. Capsules subglobose, 11-15 mm in diameter (fresh exocarp), 9-12 mm in diameter (dried endocarp). [2n=28*.] Occurring in wet forest, occasionally in divese mesic forest, 650-1,575 m, on Kaua'i. — Plate 59.

3. Trematolobelia macrostachys (Hook. & Arnott) A. Zahlbr.
[Lobelia macrostachys Hook. & Arnott; Delissea macrostachys (Hook. & Arnott) K. Presl; Dortmanna macrostachys (Hook. & Arnott) Kuntze; Trematocarpus macrostachys (Hook. & Arnott) A. Zahlbr.; Trematolobelia auriculata St. John; T. forbesii St. John; T. haleakalaensis (St. John) Degener & I. Degener; T. lustrialis St. John; T. macrostachys var. haleakalaensis St. John; T. m. var. kaalae (Degener) F. Wimmer; T. m. var. ligulata St. John; T. rockii St. John; T. sandwicensis Degener, nom. illeg.; T. s. var. kaalae Degener]
(end) Koli'i

Stems (0.5-)1-4 m long, (0.7-)1-3.5 cm in diameter. Leaves dark green on upper surface, lower surface pale green, linear-oblanceolate to narrowly elliptic, blades 9-25 cm long, 1-3.5 cm wide, glabrous, margins callose-crenulate, apex acute to acuminate, base cuneate, tapering gradually into a winged petiole 1-4.5 cm long. Inflorescences composed of 5-20 horizontal branches 20-100 cm long, pedicels 10-30 mm long, bracts narrowly oblong to lanceolate, 1-4.5 cm long, 2-6 mm wide; hypanthium 4-9 mm wide; calyx lobes erect, narrowly oblong to triangular, 3.5-15 mm long, 1-3 mm wide, apex obtuse; corolla scarlet, rose, or pink, 4-6 cm long, the tube and trifid lip gently curved downward, the 2 lateral lobes recuved, the corolla thus appearing bilabiate; staminal column 4-5.5 cm long, sparsely pubescent at apex; anthers hirsute at base. Capsules subglobose, 8-12 mm in diameter (fresh exocarp), 7-10 mm in diameter (dry endocarp). [2n=28*.] Occurring in wet forest and on windswept ridges and cliffs, in the Ko'olau (550-915 m) and Wai'anae (1,000-1,200m) mountains of O'ahu, on Moloka'i (1,200-1,500m), Lāna'i (915-975m), East Maui (915-1,300m) and West Maui (900-1,725m), and in the Kohala Mountains, Hawai'i (ca. 1,220m). — Plate 59.

page 487

page 488

4. Trematolobelia singularis St. John
(end, R)

Stems 0.6-1.5 m long, 0.6-1.5 cm in diameter. Leaves dark green on upper surface, lower surface pale green, linear-elliptic, blades 10-18 cm long, 1-1.8 cm wide, glabrous, margins callose-crenuluate, apex acute, base cuneate, tapering gradually into a winged petiole 1-2 cm long. Inflorescences unbranched, erect, 30-42 cm long, pedicels 15-35 mm long, bracts linear-lanceolate, 12-25 mm long; hypanthium 5-6 mm wide; calyx lobes erect, oblong to ovate, 4-7 mm long, 2-5 mm wide, apex obtuse; corolla violet, 4-5.5 cm long, the tube and trifid lip gently curved downward, the 2 lateral lobes recurved, the corolla thus appearing bilabiate; staminal column 32-4.5 cm long, sparsely pubescent toward apex. Capsules subglobose, 9-12 mm in diameters (fresh exocarp). Fruit and seeds unknown. [2n=28*.] Known only from a few recent specimens collected at Kōnāhuanui and Hālawa, 900-915m, on the wet windswept summit of the Ko'olau Mountains, O'ahu. — Plate 59.

9. TRIODANIS Raf.

Erect or decumbent annual herbs; stems green, 5-angled. Leaves simple, sessile to short-petiolate. Flowers sessile, usually cleistogamous, and then with vestigial corolla, stamens, and style, only the upper one chasmogamous, 1-3 in the axils of the upper leaves or arranged in terminal spike-like inflorescences; calyx 5-lobed or in cleistogamous flowers 3-4-lobed; corolla blue to purple, rotate; ovary 3-celled, placentation axile; style divided at the tip into 3 short branches. Fruit a poricidal capsule with 1 pore per cell. Seeds numerous, dark brown, smooth, lenticular, less than 1 mm long.

A genus of 8 species, one of the Mediterranean region, the remainder American. The limits and relationships of this genus are problematical (McVaugh, 1945, 1948; Fernald, 1946). It has been included by many authors in Specularia Heist. ex A. DC. Name derived from the Greek trion, three, and odontos, teeeth, in reference to the 3-lobed calyx of the cleistogamous flowers.

1. Triodanis biflora (Ruiz & Pav.) Greene
[Campanula biflora Ruiz & Pav.; Specularia biflora (Ruiz & Pav.) Fisch. & C. A. Mey.; Triodanis perfoliata (L.)Nieuwl. var. biflora (Ruiz & Pav.) Bradley]
(nat) Venus' looking-glass

Stems slender, erect or decumbent, 15-45(-80) cm long, unbranched or branched from base. Leaves broadly elliptic to ovate, 6-30 mm long, 3-12 mm wide, glabrous or hispid on the veins and margins, margins crenate, short-petiolate and obtuse below, sessile and acute above. Flowers nearly all cleistogamous, only the single terminal flower regularly chasmogamous; calyx lobes triangular to ovate, 4-7 mm long in chasmogamous flowers, 3-5 mm long in cleistogamous flowers, glabrous or sparsely pubescent; corolla tube 1-1.5 mm long, the lobes oblong, 5-9 mm long, ciliate at tips and sparsely pubescent on the veins. Capsules ellipsoid, clavate, or ovoid, 4-7 mm long, 1.3-1.7 mm wide, glabrous or sparsely pubescent, with 3 distal pores. Seeds chestnut brown, lenticular, smooth, shiny, 0.5-0.7 mm long. [2n=56.] Native to southeastern United States, northern Mexico, and temperate South America; in Hawai'i perhaps sparingly naturalized in dry, disturbed sites. the only known specimens (Fagerlund & Mitchell 1048 & 1050, BISH) were collected in Hawai'i Volcanoes National Park, Hawai'i, in 1945.

This species is very similar to Triodanis perfoliata (L.)Nieuwl., a predominantly chasmogamous species (McVaugh, 1945). Bradley (1975) has suggested that the morphological features that distinguish these 2 taxa are correlated with the observed difference in breeding system.

page 489

10. WAHLENBERGIA Schrad. ex Roth, nom. cons.

[Campanopsis (R.Br.)Kuntze, nom. illeg.; Campanula L. sect. Campanopsis R. Br.; Cervicina Delile, nom. rej.]

Erect, prostrate, or decumbent perennial herbs. Leaves pubescent or rarely glabrous, margins entire, toothed, crenate, undulate, or rarely lobed, usually cartilaginous, sessile or short-petiolate. Flowers in terminal or axillary panicles, pedicellate; hypanthium 5-nerved or 10-nerved; corolla blue to purple or white, campanulate; anthers coherent; filaments distinct; ovary inferior to rarely superior, 2-5-celled, placentation axile; style with as many lobes as cells. Fruit a hemispherical to cylindrical, apically loculicidal capsule. Seeds lenticular or trigonous, smooth or minutely reticulate.

A complex genus of approximatley 150 species, widespread in the Southern Hemisphere and extending north into Europe and Asia. The name commemorates Swedish botanist Georg Wahlenberg (1780-1851).

1. Wahlenbergia gracilis (G. Forster) A. DC
[Campanula gracilis G. Forster]
(nat)

Stems slender, angular, sometimes woody at base, (5-)15-40 cm long, unbranched to many-branched, glabrous or pubescent at base. Basal leaves rosulate and early deciduous; lower cauline leaves spatulate to obovate below, linear to lanceolate above, 1-4 cm long, 0.2-0.8 cm wide, margins entire to serrate-dentate, sessile. Calyx glabrous; corolla white to pale blue, 4-15(-25) mm in diameter; stamens dark blue to purple. Capsules obconical, 6-10 mm long. [2n=18, 36, 64.] Native to Australasia; in Hawai'i sparingly naturalized in dry, disturbed areas, locally common during wetter months, 0-2,850 m, on Moloka'i, Lāna'i, Maui, and Hawai'i. First collected in Kawela Valley, Moloka'i, in 1928 (Degener 9893, BISH). — Plate 59.

This species is extremely polymorphic, both in its original range and in Hawai'i. It has sometimes been regarded as conspecific with the Asiatic Wahlenbergia marginata (Thunb.) A. DC (Lothian, 1947; Moeliono & Tuyn, 1960).