Pacific Science (1997), vol. 51, no. 3: 254-287 © 1997 by University of Hawai'i Press. All rights reserved A Taxonomic Revision of the Endemic Hawaiian Lysimachia (Primulaceae) Including Three New Species! KENDRICK L. MARR AND BRUCE A. BOHM2 ABSTRACT: A taxonomic revision of the endemic Lysimachia of the Hawaiian Islands was undertaken with the goal of clarifying species boundaries, especially within the L. hillebrandii/L. remyi complex of the previous taxonomic treatment. The endemic species appear to be monophyletic with Malesian affinities. The revision presented here is based upon observations of morphological characters. Sixteen species are recognized, of which three are probably extinct. Most species have narrow ecological preferences and are endemic to a single island. Species differ from each other most notably in the size, shape, and venation of the leaves; the size, shape, and pigmentation of the calyx and corolla lobes; and the presence or absence of viscid stems and leaves. Populations previously classified within L. hillebrandii or L. remyi differ in a number of characters not previously evaluated including vestiture, leaf color and venation, pedicel position and color, and calyx shape and color. Three new species, L. iniki, L. pendens, and L. scopulensis, are described. A key to species, species descriptions, and distribution maps are provided. Lysimachia, ONE OF the largest genera of the sented by the indigenous coastal Lysimachia Primulaceae, consists of approximately 180 spe­ mauritiana Lam. and subgen. Lysimachiopsis cies of upright or sprawling perennial or annual (Heller) Hand.-Mazz., which consists of species herbs, shrubs, or subshrubs. Southwest China, endemic to the Hawaiian Islands. with 122 (110 endemic) species, is the center of The endemic species are perennial, scandent diversity (Chen and Hu 1979). Species also or upright, woody shrubs with alternate leaves, occur in temperate areas of the Northern Hemi­ regularly dehiscent capsules, axillary flowers, sphere, the Southeast Asian Tropics, South filaments connate and adnate to the lower third America, Africa, and Australia (Bennell and of the corolla, basifixed anthers with lateral Hu 1983). dehiscence, and a 5-10 merous perianth. Species The infrageneric classification of the genus differ from each other in vestiture, viscidness, consists of six subgenera and 18 sections and is phyllotaxy, and the size, shape, and color of based primarily on the work of Handel-Mazzetti leaves and calyx lobes. Corollas of L. glutinosa (1928), who emphasized floral structure, partic­ Rock are white to cream, whereas they are green ularly the androecium. This classification was with a red base in L. forbesii Rock and L. kala­ modified somewhat by Chen and Hu (1979). lauensis Skottsbg. and red at the base becoming Two subgenera occur in the Hawaiian Islands: white halfway toward the tip in L. iniki Marr, subgen. Palladia (Moench) Hand.-Mazz. repre- sp. nov. Corollas of the remaining species are red. The protogynous (Marr 1995), hermaphro­ 1 Supported by operating and equipment grants from dite flowers are sweetly fragrant, especially in the Natural Sciences and Engineering Research Council of the evening. Canada to B.A.B. Financial and\or logistical support to K.L.M. was provided by the Hawaiian Plant Conservation Endemic species have tetracolporate pollen Center (HPCC), The Nature Conservancy ofHawai 'i, Halea­ grains that are the largest in the genus, especially kala National Park, the Cooperative Parks Studies Unit, Uni­ those of L. forbesii Rock (Huynh 1970, 1971). versity of British Columbia graduate student travel grant, Non-Hawaiian species have tricolporate pollen, and the American Society of Plant Taxonomists. Manuscript accepted 8 September 1996. although some species occasionally produce tet­ 2 Department of Botany, University of British Columbia, racolporate grains (Huynh 1970, 1971, Bennell Vancouver, British Columbia, V6T 1Z4, Canada. and Hu 1983). 254 A Revision of Lysimachia-MARR AND BOHM 255 Species grow from 250 to 2300 m in montane revised and expanded treatment of Wagner et al. bogs, subalpine mesic shrublands, montane dry (1990) referred many of the species of St. John and wet forests, and lowland mesic shrublands (1987) within their circumscription of either L. (as classified by Gagne and Cuddihy 1990), and hillebrandii or L. remyi. on wet vertical cliffs. Species occur on all of Heller (1897) concluded that the shrubby the larger islands except Kaho'olawe, Ni'ihau, habit and urceolate, red corollas ofthe Hawaiian and Hawai'i in scattered populations and gener­ Islands species warranted the creation of a new ally have allopatric distributions. Two excep­ genus, Lysimachiopsis Heller, to include the tions are upper Kalalau Valley on Kaua'i where endemic species alone. Knuth (1905) returned the ranges of L. glutinosa, L. kalalauensis, and them to Lysimachia, but created section Frutico­ L. scopulensis Marr, sp. nov. overlap and the sae Knuth, again to include only the endemic southern Wai 'anae Range on O'ahu where L. Hawaiian species. Handel-Mazzetti (1928) com­ hillebrandii Hook. f. ex. A. Gray and L. waia­ bined sect. Fruticosae Knuth with sect. Rosula­ naeensis St. John occur together. Populations tae Champ. (containing a single Asian species, usually consist of fewer than 100 individuals. L. alpestris Champ.) to form subgenus Lysi­ An investigation of the fertility of artificial machiopsis (Heller) Handel-Mazzetti. However, interspecific F 1 hybrids is incomplete because Handel-Mazzetti synonymized Fruticosae under of the absence of flowers on greenhouse-grown a prior name, Cilicina Klatt published in 1866. L. daphnoides (A. Gray) Hillebr., L. scopulensis This is confusing and may have been in error. Marr, sp. nov., L. pendens Marr, sp. nov., L. inik In an overview of the infrageneric classification Marr, sp. nov., and L. hillebrandii. However, F 1 Ray (1956), who revised the North American hybrids that could be produced generally had species, included only part of Cilicina in subgen. pollen stainability as high as their parents with Lysimachia. This suggests that Cilicina was a the exception of some (but not all) progeny of heterogeneous group that included non-Hawai­ crosses between L. glutinosa and L. kala­ ian species. Therefore, synonymy of Cilicina lauensis, which had very low stainabilities with Fruticosae was unwarranted because Fruti­ (Marr 1995). cosae Knuth clearly included only the endemic Chromosome numbers of n = 36 are known Hawaiian species. for Lysimachia glutinosa (mistakenly identified The unique (i.e., tetracolporate) pollen of as L. kalalauensis) and L. hillebrandii Hook. f. Hawaiian Lysimachia led Huynh (1970) to split ex A. Gray (chromosomes were observed from subgen. Lysimachiopsis along sectional lines a plant now classified as L. remyi subsp. remyi) and to elevate the sections to subgeneric status: (Carr 1978). Skottsberg (1953) reported a count sect. Fruticosae became subgen. Sandwicensia of 2n = ca. 54 for L. hillebrandii from O'ahu. Huynh and sect. Rosulatae became subgen. That count, because it is reported as an approxi­ Nullicaulis Huynh (typified by L. alpestris). mate one, should be regarded as suspect. Itseems Sandwicensia was an illegitimate name (mistak­ unlikely, though not impossible, that there would enly retained by Bennel and Hu [1983]) be more than one chromosome number among because Lysimachiopsis sensu Heller (1897) the endemic species. was already available and included only the endemic Hawaiian species. Furthermore, Chen and Hu (1979) formally transferred species in Taxonomic History sect. Rosulatae from subgen. Lysimachiopsis The first description of Lysimachia from the to subgen. Lysimachia. To summarize, the Hawaiian Islands was that of Gray (1862), fol­ endemic Hawaiian Lysimachia are properly lowed by treatments ofHillebrand (1888), Heller classified in subgen. Lysimachiopsis (Heller) (1897), and Knuth (1905). All of those works Handel-Mazzetti. provided keys and descriptions as additional spe­ The taxonomic treatment of Wagner et al. cies were discovered. St. John (1987) described (1990) recognized 10 endemic species: L. daph­ 44 new endemic species and made four new noides, L. filifolia C.N. Forbes & Lydgate, L. combinations, bringing to 54 the total number forbesii, L. glutinosa, L. hillebrandii, L. kala­ of names published, but provided no key. The lauensis, L. lydgatei Hillebr., L. maxima (R. 256 PACIFIC SCIENCE, Volume 51, July 1997 Knuth) St. John, L. remyi Hillebr., and L. venosa tions from the Wailua River indicated that plants (Wawra) St. John. Lysimachia iniki Marr, sp. there typically have leaves up to 4 mm wide. nov. and L. scopulensis Marr, sp. nov. were dis­ covered after 1990. Most ofthese taxa are clearly Geographical Affinities distinct in a number of characters and present no taxonomic difficulties. However, a number A monophyletic origin for the endemic spe­ of metric characters vary among and sometimes cies is supported by several lines of evidence. within populations of L. hillebrandii and L. The most parsimonious explanation for the pres­ remyi sensu Wagner et al. (1990), such that those ence oftetracolporate pollen in all species is that authors noted (p. 1077) that Lysimachia is it was a character that evolved once, presumably "greatly in need of careful monographic work, from an ancestor with tricolporate pollen. In especially the Lysimachia hillebrandii-L. remyi characters such as the shrubby habit and variable complex on the younger islands" (i.e., Maui and