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A new autogamous species of Schiedea is described and illustrated. It is known only from cliff habitat in rainforest on a single ridge in the Natural Area Reserve, Hanawi, East Maui. With the addition of this species there are 28 species in this endemic Hawaiian genus. The new species appears to be most closely related to Schiedea nuttallii, a species of mesic habitats on O'ahu, Moloka'i, and Maui. In June 1992 James Jacobi and Arthur Medeiros were collecting in the wet forests of Hanawi, East Maui, when they discovered a species of Schiedea Chamisso & Schlechtendal they considered to be probably new to science. It appeared similar to S. nuttallii Hooker, but close study of the herbarium specimens showed that it was a closely related new species from a much wetter habitat than S. nuttallii. In June 1995 Ken Wood and Steve Perlman revisited the population with Medeiros, W. Evanson, and R. Hobdy. They obtained seeds for greenhouse propagation at the University of California, Irvine. Studies of plants grown in the greenhouse confirmed that they represented a new species, here named Schiedea jacobii. With the addition of this species there are 28 species in this endemic Hawaiian genus. Observations in the greenhouse also showed that the new species is autogamous and that it shares several characteristics with another rainforest species in the genus, S. diffusa A. Gray. These shared characters, related to the breeding system and to seed dispersal, include anthers shedding pollen while the styles are elongated and receptive, and seeds persistent on the placenta after capsule dehiscence. The latter feature may be associated with germination of seeds in the capsule, followed by dispersal of small seedlings, as noted for S. diffusa (Wagner, unpublished obs.). Schiedea jacobii W. L. Wagner, Weller & Medeiros, sp. nov. TYPE: Cultivated from seeds at University of California, Irvine greenhouse in 1997. Source: Hawaiian Islands. Maui: East Maui, Hanawi, Kuhiwa unit, E of Hanawi Stream, between Hanawai Stream and Kuhiwa drainage [20045'N, 15607'40"W], 6340 ft., Efacing slope, ca. 20 ft. below ridge top, in Metrosideros montane wet forest, 22 June 1995, S. Perlman, K. Wood, A. Medeiros, R. Hobdy & B. Evanson 14807 (holotype, US-3351775; isotypes, BISH, CANB, CHR, K, MO, NY, PTBG). Figure 1A-H. Haec species S. nuttallii similis, sed ab ea caulibus quadrangularibus, floribus autogamis, sepalis sub angulo ca. 100-135 e pedicello abeuntibus, tubo apicali 1.4-1.8 Figure 1. Schiedea jacobii W. L. Wagner, Weller & Medeiros. -A. Flowering stem. -B. Stem cross section. -C. Portion of inflorescence branch. -D. Flower, showing stamens ready to dehisce and styles elongated and receptive. -E. Adaxial view of sepal with nectary and stamen. -F. Dehisced capsule surrounded by the sepals exposing the mass of persistent seeds. -G. Seed, lateral view. -H. Seed, marginal view. Drawn from live greenhouse material of the type. NovoN 9: 284-287. 1999. This content downloaded from 157.55.39.136 on Thu, 19 May 2016 04:39:19 UTC All use subject to http://about.jstor.org/terms Volume 9, Number 2 1999 Wagner et al. Schiedea jacobii 285

KANEHIRO KITAYAMA*, EDWARD A.G. SCHUURt, DONALD R. DRAKE: and DIETER MUELLER-DOMBOIS? *The Japanese Forestry and Forest Products Research Institute, PO Box 16, Tsukuba Norin Kenkyu Danchi, Ibaraki 305, Japan, tEcosystem Sciences Division, University of California at Berkeley, Berkeley, California 94720, USA, tSchool of Biological Sciences, Victoria University of Wellington, PO Box 600, Wellington, New Zealand, and ?Department of Botany, University of Hawaii, 3190 Maile Way no. 101, Honolulu, Hawaii 96822, USA

A new species in the Lycopodiaceae, a firmoss, Phlegmariurus stemmermannii A.C. Madeiros & W.H.Wagner, is described and illustrated from East Maui and the island of Hawaii. A contrasting description is also given for the most closely related Hawaiian species, P. mannii (Hillebr.) W.H.Wagner. The non-strobilate Lycopodiaceae in Hawaii comprise four species and one hybrid of epiphytic, non-gemmiferous firmosses (Phlegmariurus) and five species and five hybrids of terrestrial or epipetric, gemmiferous firmosses (Huperzia). Of the former, only one of the taxa (Phlegmariurus filiformis (Sw.) W.H. Wagner) lacks pendent tassles of sporophylls, and the remainder possess the tassles. The rarest of the tassled firmosses is Mann's firmoss (P mannii). Very few collections have been made of it, all from the mountains of West and East Maui, and Hawaii. It has been studied intensively in the field by Medeiros and Hobdy over the past decade, and there seems to be no question regarding its species status. During studies of the collections at the Bishop Museum in 1993, however, W. H. Wagner recognized an unusual single specimen from Hawaii that appeared to be a new species. Subsequently, plants with the same characteristics were discovered by Medeiros in Haleakala, in East Maui. With these additional specimens it was possible to confirm the distinctive features of the new lycopod, and the results of these studies are reported here. The two species dealt with in this paper belong to 11gaard's Myrsinites subgroup, "Slender to delicate plants, often with red stems, usually decussate or subdecussate leaves throughout. Leaves of the basal divisions irregular in size or shape, usually not overlapping continuously" (0llgaard 1987), with approximately a dozen species, all in tropical America, except for those in Hawaii. The Hawaiian species may be recognized by the fact that the sterile portions of the shoots are relatively short, usually considerably less than 15 cm, and the fertile or partially fertile tassles are usually long, up to 25 cm, Author for correspondence. This content downloaded from 157.55.39.237 on Thu, 07 Jul 2016 05:04:56 UTC All use subject to http://about.jstor.org/terms AMERICAN FERN JOURNAL: VOLUME 86 NUMBER 3 (1996) except in young plants. They grow on rough-barked trees, mostly Acacia koa, 3 meters or less from the ground. The generic names used here are those discussed by Wagner and Beitel (1992). The following key separates these two species (see also Table 1): Fertile tassles branching angles mostly 70?-140?; ultimate fertile branches 1.0-1.5 mm thick; sporophylls broadly rounded to slightly pointed, mostly short-ovate, 1-2 mm long; sterile basal portions of stem bright red 1. Phlegmariurus mannii Fertile tassles branching angles mostly 10?-50?; ultimate fertile branches 1.5-2.2 mm thick; sporophylls more pointed, mostly triangular, 1.5-2.5 mm long; sterile basal portions of stems green to pale yellow 2. Phlegmariurus stemmermanniae 1. Phlegmariurus mannii (Hillebr.) W.H. Wagner in W.H. Wagner, Jr., F.S. Wagner, & T. Flynn, Contr. Univ. Michigan Herb. 20:241. 1995 (Figs. 1, 2).Lycopodium phlegmaria L. var. mannii Hillebr., Fl. Hawaiian Isl. 645. 1888.-Lycopodium mannii (Hillebr.) Skottsb.-TYPE: Hawaii, West Maui, mountains above Malaea Bay, Mann & Brigham 506 (CU; isotype BISH). Pendent epiphyte on bark of trees. Sterile basal stem red. Intervals between basal sterile leaves along a row (typical trophophylls) remote, 2-9 mm apart. Sterile leaf tip indurated, apiculum mostly sharp, 0.2-0.3 mm long. Sterile to fertile branch transition mostly abrupt. Length of fertile, terminal branchlets 0.5-8 cm (mostly less than 3 cm). Maximum number of fertile terminal branches for a whole shoot 16-45. Branching angle of first 0.5 cm of fertile branches spreading widely, 70?-140?. Basal 1.5-3.5 cm of fertile branches with pronounced curvature, becoming straighter and more parallel distally. Sporophylls (with visible sporangia) 1.0-2.0 mm long, mostly rounded apically to slightly pointed, equal to or slightly shorter than exposed sporangia. Thickness of branches (with visible sporangia) 0.8-1.1 mm. Other Collections: HAWAII. Hawaii. NW of Papaloa, Captain Cook, Kona, epiphytic on giant koas at 1525 m, Degener 20330 & Murashigu (BISH-2 sheets). East Maui. Above W Rim of Manawainui Canyon Kaupo Maui, crotch of Acacia koa tree 2.5 m above ground, many young plants growing together to form a dense mat 0.6 m across, sporophylls sometimes root when touching suitable sites, 1300 m, Hobdy 1178 (BISH); Kahikinui, Manawainui Gulch, in rough scaly bark of an old Acacia koa tree in a rich native forest remnant, 1.5 m above the ground, 1600 m, Hobdy 2273 (MICH-2 sheets); S slope of Haleakala, Manawainui Gulch, 6 plants seen growing on two Koa trees, 1600 m, Medeiros 310 (BISH); W side of Nuanaloa Stream, Kipahulu, vegetation: Athyrium, Sadleria, Acacia, Cheirodendron, and Metrosideros, 1300 m, Higashino & Mizuno 3140 (BISH). West Maui. Haelaau, on tree trunk, 885 m, Ewart & Swezey 135 (BISH, US, UC); Lihau Peak, epiphytic on ohia tree in fairly dense shady understory 3 m above ground, 1200 m, Hobdy 22422 (BISH). 2. Phlegmariurus stemmermanniae A.C. Medeiros & W.H. Wagner, sp. nov. (Figs. 3, 4).-TYPE: Hawaii, Maui, SE slope of Haleakala, Kaapahu, epiphytic on Syzygium sandwicense tree, 1150 m, 1995, Medeiros & Chimera 2220 (MICH). Differt a Phlegmariuro mannii caulibus sterilibus omnibus viridibus ad caules fertiles gradatim mutantibus, foliis confertioribus 1-2 mm inter se distan90 This content downloaded from 157.55.39.237 on Thu, 07 Jul 2016 05:04:56 UTC All use subject to http://about.jstor.org/terms MEDEIROS ET AL.: A NEW HAWAIIAN PHLEGMARIURUS

Metrosideros polymorpha Gaud. (MYRTACEAE), 'ohi'a, dominates pioneer dry forest communities in Hawai'i, while other tree taxa dominate later seral stages. Tree density is greater in the later seral communities, and the dominant species there exhibit significantly lower mid-day plant water potentials than Metrosideros from nearby pioneer communities. This pattern suggests that the greater water stress experienced by the later seral trees may be due to competition for limited water, and this may restrict Metrosideros from these sites. The replacement of Metrosideros in later seral stages of Hawaiian dry forests contrasts with its maintenance as the dominant canopy component in both early and late seral rain forest communities.

The vegetation of the Puhimau geothermal area was compared with that of the surrounding ecosystem. Very few species of the surrounding area were found with any abundance in the geothermal area. The center of the geothermal area is almost barren. Pockets of vegetation dominated by the endemic moss Campylopus praemorsus (C. Muell.) Jaeg. are scattered throughout the central area. The only other temperature-tolerant species present is Portulaca sclerocarpa Gray. Cladonia oceanica Vain. is commonly found growing epiphytically on the moss away from the high temperatures. The central zone is surrounded by an Andropogon-dominated grassland in which both C. praemorsus and C. oceanica are commonly fertile. No lichens exhibited any tolerance of high temperatures. The Puhimau geothermal area, often referred to locally as the Puhimau Hot Spot, is located on the east rift zone of Kilauea volcano on the island of Hawaii. The area was formed in the winter of 1937-38 when magma came close to the surface and resulted in hydrothermal activity rather than the usual destruction associated with lava flows. The original Metrosideros forest was killed but not burned. The steam vents formed were confined principally to the bases of tree trunks. Some of these trunks are still present in the area and support a distinctive fumarolic community. Although sulphurous gases are normally emitted from fumaroles during the degassing of subterranean lava, no sulphur dioxide or hydrogen sulphide gas has been recorded from the vents in the Puhimau thermal area in recent years (Siegel & Siegel 1979). Recent subterranean volcanic activity in the area resulted in a short (less than 48 hours) emission of SO2. Fosberg and Lamoureux (1966) briefly described the vegetation of two areas of the east rift zone of Kilauea in Hawaii Volcanoes National Park in which there was hydrothermal activity. One of these areas has since been destroyed in the formation of Mauna Ulu. In these areas, they recorded Cladonia cf. cristatella (= C. oceanica Vain.), C. cf. rangi- ferina (= C. skottsbergii Magn.), and Racomitrium (presumably R. lanuginosum var. pruinosum Wils.). Kappen (1973), citing a personal communication from Mueller-Dom- bois, confirmed the presence of these lichens "scattered on volcanic soils." Ground tem- peratures from 48 to 89?C have been recorded in both areas, suggesting that all species growing there were remarkably heat tolerant. Although lichens (Lange 1953; Kappen 1973) and mosses (Noerr 1974) are very tolerant of high temperatures when desiccated, when hydrated their tolerance is significantly di- minished. Some mosses are tolerant of temperatures up to 50'C (Noerr 1974) whereas lichens can remain active only up to about 25-30?C (Lange 1965).

Myrtus metrosideros was described by Bailey based on a leaf collection fiom Mt Bellenden Ker in northern Queensland. It was so unlike other species of Myrtus that Diels (1922) considered that it possibly belonged to his new genus Xanthomyrtus. When Burret (1941) transferred the Australian species of Myrtus to Austromyrtus he made the combination for M. metrosideros based on the description only. In the same article, Burret described the genus Uromyrtus with six to nine species in New Caledonia. In 1979 I transferred three species to Uromyrtus and described three new species, including U. australis the first record of Uromyrtus from Australia. At that time I had only seen leaf collections of M. metrosideros, but since then I have examined collections with both flowers and fruits that match the type collection; these possess the anther appendages that distinguish Uromyrtus from Myrtus. The new combination is therefore made recognising the second species of Uromyrtus from Australia.

Xanthomyrtus was described by Diels (1922) with thirteen species in New Guinea, one in Sabah and one in New Caledonia. In addition Myrtus metrosideros Bailey, from northern Queensland, was suggested by Diels as possibly belonging to Xanthomyrtus. Diels distinguished Xanthomyrtus from Myrtus and related genera by the 4-merous flowers with thick triangular calyx lobes, the yellow, clawed petals, the few (I-2-)-seriate stamens and the 3-flowered dichasia with sessile flowers. Diels described eight new species of Xanthomyrtus, the remaining species were transferred by him from Eugenia or Myrtus; three of these were described by Ridley (1916) from West New Guinea, three described by Gibbs (1917) from the Arfak Mountains and one based on Myrtusflavida Stapf from Borneo. Diels (1924) later added one more species from New Guinea and cited further collections. However, in some of the species that he transferred to Xanthomyrtus the original collections had not been seen by him and were only known to him by the descriptions or by collections matched with descriptions. Many species were known only from a single or a few specimens and Diels (1922) cited only 24 collections. Merrill (1926) transferred Eugenia aurea Elmer and E. diplicosifolia C. B. Robinson, both from the Philippines, to Xanthomyrtus and later (1928) transferred two further Bornean species, Myrtus taxifolia Ridl. and M. moultonii Merr. Guillaumin (i939) recognized two species in New Caledonia, X. hienghenensis Guillaumin, with one variety, and X. sarasinii (Guillaumin) Guillaumin, including X. pergracilis Diels as a synonym of the latter. Merrill and Perry (1942) reviewed the species of Xanthomyrtus in New Guinea and described seven new species. Unfortunately many of the species previously recognized by Diels were known to them only through descriptions and specimens matched with the descriptions; several of the species described by them were incompletely known and they cite only 37 collections. Myrtus metrosideros, which tentatively was included in Xanthomyrtus by Diels (1922), is based on a leaf collection from northern Queensland. However, further collections matched with the type belong to Myrtus sect. Austromyrtus and no collections of Xanthomyrtus have been seen from Australia, although the genus may be expected to occur there. Xanthomyrtus pergracilis has been examined and is a species of Eugenia sens. str., and only a single species, X. hienghenensis, is recognized from New Caledonia; this species appears to be closely related to species in New Guinea.