Merger of the Endemic Hawaiian Genera Cyanea and Rollandia (Campanulaceae: Lobelioideae)

5 Pages·1993·3.2 MB·English

by  Thomas G Lammers

Merger of the Endemic Hawaiian Genera Cyanea and Rollandia
(Campanulaceae: Lobelioideae)
Th omas G. hammers
Department of Botany, Field Museum of Natural History, Chicago,
Illinois 60605-2496, U.S.A.
Thomas J. Givnish and Kenneth J. Sytsma
Department of Botany, University of Wisconsin, Madison, Wisconsin 53706-1381, U.S.A.
ABSTRACT. Phylogenetic analyses based on mo¬ Rollandia as distinct genera. The traditional four-
lecular data indicate that the species of Rollandia genus classification was employed in the most recent
are embedded within the genus Cyanea and that taxonomic summary of the Hawaiian Campanula¬
together these genera form a inonophyletic group. ceae (Lammers, 1990).
Cyanea and Rollandia are therefore merged under Recent phylogenetic analyses based on restriction
the former name. A revised key to the baccate site variation in the chloroplast DNA of Hawaiian
genera of Hawaiian Fobelioideae and an expanded lobelioids (Givnish et al., in press) have shown that
description of the genus are provided; eight new' the species comprising Rollandia are embedded
combinations and one new name are proposed; and within Cyanea as traditionally circumscribed, that
lectotypes are selected for Rollandia humboldtiana Cyanea without Rollandia is paraphyletic, and that
and R. longi flora var. angustifolia. This expanded the two genera together form a monophyletic group.
circumscription makes Cyanea the largest genus in Specifically, Rollandia is part of a major clade that
the Hawaiian flora, with 64 species and 10 heter- is defined by 10 synapomorphic mutations and gen¬
onymic subspecies. erally characterized by orange fruits and an un¬
branched habit. Within this major clade, Rollandia
The flora of the Hawaiian Islands includes 114 forms an unresolved clade defined by three muta¬
endemic species of woody Lobelioideae (Campanu¬ tions.
laceae), 95 of which have baccate fruits (hammers, Morphologically, the two genera are very similar.
1990, 1992a; Lammers & Lorence, 1993). Tra¬ Every author for over 100 years has distinguished
ditionally (Gaudichaud, 1826; Bentham & Hooker, Rollandia solely on the basis of a single feature,
1876; Hillebrand, 1888; Schonland, 1889; Hock, i.e., the adnation of the stamina! column to the
1919; Wimmer, 1943), these baccate species have corolla (Hillebrand, 1888; Schonland, 1889; von
been divided among four genera: Clermontia Gau¬ Post & Kuntze, 1903; Hock, 1919; Wimmer, 1943;
dichaud, Cyanea Gaudichaud, Delissea Gaudi¬ Lammers, 1990). The traditional classification of
chaud, and Rollandia Gaudichaud. However, var¬ Campanulaceae includes several such cases of close¬
ious alternative classifications have been proposed. ly related genera distinguished by a single morpho¬
Presl (1836) and de Candolle (1839) recognized a logical feature, e.g., Campanumoea Blume and Co-
fifth genus, Macrochilus C. Presl. Gray (1861), donopsis Wallich, Lobelia and Pratia Gaudichaud.
however, recognized only three; Macrochilus was The circumscription of such genera must be revised
included in Cyanea and the species of Rollandia if a natural classification of the family is ever to be
were divided between Cyanea and Delissea. En- achieved (Lammers, 1992b, c).
dlicher (1836) and Dietrich (1839), taking a very In order to provide a more natural classification
broad view of generic circumscription, assigned all of the baccate Hawaiian lobelioids, Cyanea and Rol¬
Hawaiian lobelioids to cosmopolitan Lobelia L. Bail- landia are here merged. Although Gray (1861)
Ion (1885), on the other hand, treated the baccate transferred some species of Rollandia to Cyanea.
Hawaiian species as a single genus, Delissea, with the species that was later designated as the lectotype
the four traditional genera as sections. The classi¬ of the genus (R. lanceolata) was assigned to De¬
fication of von Post and Kuntze (1903) was similar, lissea. Because no previous author has effectively
except Rollandia was maintained as distinct. St. merged just Cyanea and Rollandia, and because
John (1987) merged Cyanea and Delissea under the two generic names are of equal priority, we have
the latter name, but maintained Clermontia and the prerogative of choosing the name to be retained.
Novon 3: 437-441. 1993.
438 Novon
We here choose Cyanea, as doing so will require long, white, dull, conspicuously transverse¬
ly rugose.Delissea
far fewer nomenclatural innovations.
2b. Corolla smooth or rarely muricate, without
The eight species of Rollandia (one comprising
definite dorsal or lateral knobs; seeds 0.2
two subspecies) recognized by hammers (1990) are 0.5(—2) mm long, dark brown to black,
here transferred to Cyanea with no change in their shiny, smooth or microscopically foveate-
circumscription or rank. In addition, lectotypes are reticulate. Cyanea
designated for two inadequately typified names. De¬
scriptions, distributional and habitat data, and lists Cyanea Gaudichaud, Voy. Uranie 457. 1829. De¬
of heterotypic synonyms are found in hammers’s lissea sect. Cyanea (Gaudichaud) Baillon, Hist.
(1990) treatment and will be amplified in the forth¬ PI. 8: 364. 1885. Kittelia Ih G. h. Reichen-
coming monograph of Cyanea (hammers, in prep.). bach, Handb. Nat. Pfl.-Syst. 186. 1837, nom.
Assignment of these species to an infrageneric taxon illeg. TYPE: Cyanea grimesiana Gaudichaud.
within Cyanea is problematic. As noted previously
Rollandia Gaudichaud, Voy. Uranie 458. 1829, syn.
(hammers, 1990), the infrageneric classification of nov. Delissea sect. Rollandia (Gaudichaud) Baillon,
Cyanea is badly in need of revision. The genus has Hist. PI. 8: 364. 1885. TYPE: Rollandia lanceo-
been divided into either three (Hillebrand, 1888; lata Gaudichaud (selected by Rock (1919: 363)).
Macrochilus C. Presl, Prodr. Monogr. Lobel. 47. 1836.
Wimmer, 1943; St. John, 1969) or five sections
TYPE: Macrochilus superbus (Chamisso) C. Presl
(Rock, 1919; Stone, 1967). Neither classification (based on Lobelia superba Chamisso).
maps cleanly onto the molecular clades defined by
Givnish et al. (in press); for example, Cyanea sect. Shrubs, treelets, or trees, rarely lianas, terrestrial
Palmiformes (Hillebrand) Rock comprises species or rarely epiphytic; stems unbranched or sparingly
branched, erect or ascending, unarmed, muricate,
belonging to at least three different clades. A definite
infrageneric assignment is deferred, therefore, pend¬ or aculeate (more so in juveniles), with numerous
ing completion of a comprehensive reevaluation of helically arranged leaf scars near apex; pith solid
relationships within the enlarged Cyanea. or chambered; latex white, yellow, or tan, viscous.
This merger makes Cyanea the largest genus in Leaves alternate, petiolate or sessile, glabrous or
pubescent, sometimes muricate or aculeate, es¬
the Hawaiian flora, with 64 species and 10 heter-
pecially on the petiole; margin entire, callose-toothed,
onymic subspecies (Wagner et ah, 1990; hammers
or pinnately lobed, cleft, parted, or divided (more
& horence, 1993). A morphological description of
so in juveniles), the lobes or segments then entire,
the newly enlarged genus is provided below, as is a
callose-toothed, lobed, or divided. Inflorescences ax¬
revised key to all of the Hawaiian genera of hobe-
illary, racemose, (3-)5-25(-40)-flowered, the ra-
lioideae with baccate fruit. Both are adapted from
chis sometimes obsolete and the raceme thus ap¬
hammers (1990).
pearing subumbellate. Flowers epigynous, resupinate,
A brief comment on the spelling of Hawaiian place
protandrous. Calyx synsepalous; tube adnate to the
names in the specimen citations is in order. The
ovary, forming a hypanthium, obconic or obovoid,
spellings adopted here are those given by Pukui et
rarely ovoid, oblong, cylindrical, or depressed ob¬
ah (1974), in which the macron (—) is used to indicate
ovoid; lobes 5, valvate, persistent, dentiform, tri¬
long vowels and the hamza (4) to indicate glottal
angular, or oblong, rarely linear, distinct or rarely
stops. Although non-Polynesian authors are often
connate, much shorter to somewhat longer that the
tempted to omit these diacritical marks, doing so is
hypanthium, rarely as long as the corolla. Corolla
a grammatical gaffe comparable to omitting the um¬
zygomorphic, bilabiate or unilabiate, white or various
lauts from German words or the tildes from Spanish
shades of magenta, purple, or pink, rarely greenish
ones.
or yellowish, sometimes longitudinally striped, gla¬
brous or pubescent, rarely muricate; tube suberect,
Revised Key to the Baccate Hawaiian Genera of Lobe- curved, arcuate, or sigmoid, dorsally cleft to about
LIOIDEAE the middle; lobes 5, valvate, spreading, deflexed, or
rarely erect. Stamens 5, syngenesious, included or
la. Stems repeatedly branched, spreading; inflores¬
cences (l-)2-3(-10)-flowered; corolla cleft dor- exserted; filaments connate, free from the corolla
sally nearly to base .Clermontia or dorsally adnate for %-% its length; anthers con¬
lb. Steins unbranched or only sparingly branched, nate, dithecal, opening introrsely by longitudinal slits,
erect or ascending; inflorescences (3—)5—25 the 3 dorsal ones a little longer than the 2 ventral
( 40)-flowered; corolla cleft to about the middle.
ones, the latter or rarely all 5 with apical tufts of
2a. Corolla with a small knob at the terminus
of the dorsal cleft, sometimes with two ad¬ white hairs; pollen tricolporate, prolate, ellipsoidal.
ditional lateral knobs; seeds 0.7-1.5 mm Ovary 2-loculed; placentae large, axile; style slen-
Volume 3, Number 4 Lammers et al. 439
1993 Merger of Cyanea and Rollandia
der, terete, with a ring of stiff white hairs near the Only one other syntype, a sheet at GH labeled
apex; stigma 2-lobed, the lobes appressed and non- “Oahu from Manoa Eastward,” has been seen by
receptive as the style grows through the anther tube, Lammers. A specimen at B cited by Rock (1918,
pushing out pollen, after which the stigmas spread 1919) could not be located; presumably, it was among
and become receptive. Fruit a yellow, orange, or those destroyed when the herbarium burned in 1943
purple berry. Seeds numerous, small, dark brown (Hiepko, 1987).
to black, shiny, smooth or microscopically foveate-
reticulate. Chromosome number n = 14 (Lammers, Cyanea lanceolata (Gaudichaud) Lammers, Giv¬
1988). nish & Sytsma, comb. nov. Basionym: Rollan¬
dia lanceolata Gaudichaud, Voy. Uranie 458.
1829. Lobelia lanceolata (Gaudichaud) Hook¬
Cyanea crispa (Gaudichaud) Lammers, Givnish &
er & Arnott, Bot. Beechey Voy. 88. 1832; not
Sytsma, comb. nov. Basionym: Rollandia cris¬
Hooker & Arnott, Bot. Beechey Voy. 301.
pa Gaudichaud, Voy. Uranie 459. 1829. Lo¬
1838; nor Kuntze, Revis. Gen. PI. 378. 1891.
belia crispa (Gaudichaud) Endlicher, Ann.
Lobelia rollandia Gaudichaud ex I). Dietrich,
Wiener Mus. Naturgesch. 1: 170. 1836; not
Syn. PI. 1: 728. 1839, nom. illeg. Delissea
Graham, Edinburgh New Philos. J. 1: 173.
lanceolata (Gaudichaud) A. Gray, Proc. Amer.
1826. Cyanea rollandia A. Gray, Proc. Amer.
Acad. Arts 5: 147. 1861. TYPE: “lies Sand¬
Acad. Arts 5: 149. 1861, nom. illeg. TYPE:
wich.,” Gaudichaud s.n. (holotype, P; isotype,
‘Ties Sandwich.,” Gaudichaud s.n. (holotype,
G).
P).
Based on Gaudichaud’s itinerary (St. John & Tit-
Based on Gaudichaud’s itinerary (St. John & Fit-
comb, 1983) and the known range of the species
comb, 1983) and the known range of the species
(Lammers, 1990), the type probably was collected
(Lammers, 1990), the holotype probably was col¬
on 0‘ahu, in the Ko‘olau Mountains above Honolulu,
lected on 0‘ahu, in the Ko‘olau Mountains above
during 26-29 August 1819.
Honolulu, during 26 29 August 1819.
Cyanea huinboldtiana (Gaudichaud) Lammers, Cyanea lanceolata subsp. calycina (Chamisso)
Givnish & Sytsma, comb. nov. Basionym: Rol¬ Lammers, Givnish & Sytsma, comb. nov. Bas¬
landia humboldtiana Gaudichaud, Voy. Bon- ionym: Lobelia calycina Chamisso, Linnaea 8:
ite, Bot., pi. 76. 1842. TYPE: “lies Sand¬ 222. 1833. Rollandia calycina (Chamisso) G.
wich.,” Oct. 1836, Gaudichaud s.n. (lectotype, Don, Gen. Hist. 3: 699. 1834. Delissea ca¬
selected here, W). lycina (Chamisso) C. Presl, Prodr. Monogr.
Lobel. 47. 1836. Rollandia lanceolata suhsp.
Both Rock (1919) and St. John (1940) failed to
locate type material of this species among Gaudi¬ calycina (Chamisso) Lammers, Syst. Bot. 13:
chaud’s specimens at P; Lammers likewise was un¬ 507. 1988. TYPE: Hawaii. “0 Wahu” [i.e.,
0‘ahu]: Chamisso s.n. (holotype, LE).
able to find any during a visit in September 1987.
However, he did discover one sheet ot original ma¬ Cyanea longiflora (Wawra) Lammers, Givnish &
terial at W, which is here designated as the lecto¬ Sytsma, comb. nov. Basionym: Rollandia lon¬
type. Based on Gaudichaud’s itinerary (Lasegue, giflora Wawra, Flora 56: 44. 1873. TYPE:
1845) and the known range of the species (Lam¬
Hawaii. 0‘ahu: leg. Hillebrand, If awra 2285
mers, 1990), the type probably was collected on (holotype, W).
O'ahu, in the Ko‘olau Mountains above Honolulu.
Cyanea parvifolia (C. Forbes) Lammers, Givnish
& Sytsma, comb. nov. Basionym: Rollandia
Cyanea koolauensis Lammers, Givnish & Syts¬
parvifolia C. F orbes, Occas. Pap. Bernice Pau-
ma, nom. nov. Replaced name: Rollandia an-
ahi Bishop Mus. 5: 10. 1911. TYPE: Hawaii.
gustifolia (Hillebrand) Rock, Bull. Torrey Bot.
Kaua‘i: Wai‘oli Valley, 1 Aug. 1909, Forbes
Club 45: 136. 1918; not Cyanea angustifolia
103.K (holotype, BISH).
(Chamisso) Hillebrand, FI. Hawaiian Isl. 253.
1888. Basionym of replaced name: Rollandia Cyanea purpurellifolia (Rock) Lammers, Giv¬
longiflora var. angustifolia Hillebrand, FI. Ha¬ nish & Sytsma, comb. nov. Basionym: Rollan¬
waiian Isl. 246. 1888. TYPE: Hawaii. O'ahu: dia purpurellifolia Rock, Coll. Hawaii Publ.
Palolo Valley, Hillebrand s.n. (lectotype, se¬ Bull. 2: 44. 1913. TYPE: Hawaii. 0‘ahu:
lected here, BISH). Hau‘ula ridge, Punalu‘u Mts., 2,000 ft., Aug.
440 Novon
1911, Rock 8821 (holotype, BISH; isotypes, W. L. Wagner & V. A. Funk (editors), Patterns of
BISH, GH, K, W). Speciation and Biogeography in the Hawaiian Biota.
Smithsonian Institution Press, Washington.
Cyanea st.-johnii (llosaka) Lammers, Givnish & Gray, A. 1861. Notes on Lobeliaceae, Goodeniaceae,
Sytsma, comb. nov. Basionym: Rollandia st.- &c. of the collections of the U.S. South Pacific Ex¬
ploring Expedition. Proc. Amer. Acad. Arts 5: 146-
johnii Hosaka, Occas. Pap. Bernice Pauahi
152.
Bishop Mus. 11(13): 14. 1935. TYPE: Hawaii.
Hiepko, P. 1987. The collections of the Botanical Mu¬
0‘ahu: Ko‘olau Range, Waipi‘o, Klpapa Gulch, seum Berlin-Dahlem (B) and their history. Englera
wet windswept main divide, 2,800 ft., 6 Aug. 7: 219-252.
1933, Hosaka 1159 (holotype, BISH). Hillebrand, W. 1888. Flora of the Hawaiian Islands:
A Description of their Phanerogams and Vascular
Cryptogams. Williams & Norgate, London.
Acknowledgments. Fieldwork by TGL was suit-
Lammers, T. G. 1988. Chromosome numbers and their
ported by National Science Foundation (NSF) Dis¬
systematic implications in Hawaiian Lobelioideae
sertation Research Grant BSR-8313285 (T. F. (Campanulaceae). Amer. J. Bot. 75: 1130-1134.
Stuessy, principal investigator) and a Sigma \i Grant- -. 1990. Campanulaceae. Pp. 420-489 in W.
in-Aid of Research. Travel by TGL to American and L. Wagner, D. R. Herbst & S. H. Sohmer, Manual
of the Flowering Plants of Hawai‘i. Univ. Hawai'i
European herbaria, respectively, was made possible
Press, Honolulu.
by an Herbarium Travel Award from the American -. 1992a. Two new combinations in the endemic
Society of Plant Taxonomists and a Graduate Stu¬ Hawaiian genus Cyanea (Campanulaceae: Lobelioi¬
dent Alumni Research Award from The Ohio State deae). Novon 2: 129-131.
-. 1992b. New combinations for Asian Cam¬
University. The staff's of B, BISH, G, GH, k. LE,
panulaceae. Bot. Bull. Acad. Sin. 33: 285-287.
P, PTBG, and W are acknowledged for the loan of
-. 1992c. Systematics and biogeography of the
specimens and for courtesy extended to TGL during Campanulaceae of Taiwan. Pp. 43-61 in C.-I Peng
visits to their institutions. (editor). Phytogeography and Botanical Inventory of
Field studies and molecular systematic analyses Taiwan. Institute of Botany, Academia Sinica, Taipei.
- & D. H. Lorence. 1993. A new species of
by IJG and KJS were supported by a grant from
Cyanea (Campanulaceae: Lobelioideae) from Kaua'i,
the National Geographic Society, and by NSF grants
and the resurrection of C. remyi, Novon 3: 431-
BSR-9007293 and BSR-9020055; additional logis¬ 436.
tic support was provided by the National Tropical Lasegue, A. 1845. Musee Botanique de M. Benjamin
Botanical Garden. TJG and KJS extend their heart¬ Delessert. Notices sur les Collections de Plantes et
la Bibliotheque. Librairie de Fortin, Masson et Cie.,
felt thanks to S. Anderson, L. Cuddihy, T. Flynn,
Paris.
R. Hobdy, J. Jacobi, I). Lorence, A. Medeiros, E. Presl, C. B. 1836. Prodromus Monographiae Lobeli-
Misaki, J. Obata, S. Perlman, and K. Woods, who acearum. Theophilus Haase, Prague.
accompanied them in the field and helped locate Pukui, M. K., S. H. Elbert & E. T. Mookini. 1974.
Place Names of Hawaii, rev. ed. Univ. Hawai'i Press,
many rare species of Hawaiian lobelioids.
Honolulu.
Rock, J. F. 1918. New species of Hawaiian plants. Bull.
Literature Cited
Torrey Bot. Club 45: 133-139 + pi. 6.
Baillon, H. 1885. Lobelieae. Pp. 362-367 in Histoire -. 1919. A monographic study of the Hawaiian
des Plantes, vol. 8. Librairie Hatchette et Cie., Paris. species of the tribe Lobelioideae family Campanula¬
Bentham, G. & J. D. Hooker. 1876. Genera Plantarum, ceae. Mem. Bernice Pauahi Bishop Mus. 7(2): i-xvi,
vol. 2, part 2. Reeve, London. 1-395.
Candolle, A. L. P. P. de. 1839. Lobeliaceae. Pp. 339- St. John, H. 1940. Ophioglossum, Rollandia, and
413, 784-786 in A. P. de Candolle, Prodromus Scaevola. Hawaiian plant studies 9. Occas. Pap.
Systematis Naturalis Regni Vegetabilis, vol. 7. Treut- Bernice Pauahi Bishop Mus. 15: 351-359.
tel & Wiirtz, Paris. -. 1969. Types of sections in Clermontia, Cy¬
Dietrich, D. 1839. Synopsis Plantarum, vol. 1. Voigt, anea, and Delissea (Lobeliaceae). Taxon 18: 483.
Weimar. -. 1987. Enlargement of Delissea (Lobeliaceae).
Endlicher, S. 1836. Bemerkungen fiber die Flora der Hawaiian plant studies 138. Phytologia 63: 79-90.
Sfidseeinseln. Ann. Wiener Mus. Naturgesch. 1: 129 - & M. Titcomb. 1983. The vegetation of the
190. Sandwich Islands as seen by Charles Gaudichaud in
Gaudichaud, C. 1826. Botanique. In L. de Freycinet, 1819. A translation, with notes, of Gaudichaud’s
Voyage autour du Monde, Entrepris par Ordre du “lies Sandwich." Bishop Mus. Occas. Pap. 25(9):
Roi . . . Execute sur les Corvettes de S. M. l’Uranie 1-16.
et la Physicienne, pendant les Annees 1817, 1818, Schonland, S. 1889. Campanulaceae. Pp. 40-70 in A.
1819 et 1820. Pillet-aine, Paris. Engler, Die Natfirlichen Pflanzenfamilien, Teil IV,
Givnish, T. J., K. J. Sytsma, J. F. Smith & W. J. Hahn. Abteilung 5. Wilhelm Engelmann, Leipzig.
In press. Molecular evolution, adaptive radiation, Stone, B. C. 1967. A review of the endemic genera of
and geographic speciation in Cyanea (Campanula- Hawaiian plants. Bot. Rev. 33: 216-259.
ceae), the largest plant genus endemic to Hawai'i. In von Post, T. & O. Kuntze. 1903. Lexikon Generum
Volume 3, Number 4 Lammers et al. 441
1993 Merger of Cyanea and Rollandia
Phanerogamarum. Deutsche Verlags-Anstalt, Stutt- Wimmer, F. E. 1943. Campanulaceae-Lobelioideae. I.
gart. Teil. Pp. i-vi, 1-260 in R. Mansfeld (editor). Das
Wagner, W. I,., D. R. Herbst & S. H. Sohrrier. 1990. Pflanzenreich, Teil IV, Abteilung 276b. Wilhelm En-
Manual of the Flowering Plants of Hawai'i. Univ. gelmann, Leipzig.
Hawai'i Press, Honolulu.