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A Revision of the Genus Sadleria (Blechnaceae)L

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A Revision of the Genus Sadleria (Blechnaceae)L Pacific Science (1997), vol. 51, no. 3: 288-305 © 1997 by University of Hawai'i Press. All rights reserved A Revision of the Genus Sadleria (Blechnaceae)l DANIEL D. PALMER2 ABSTRACT: The genus Sadieria is revised. Problems with nomenclature and species descriptions are reviewed and clarified. New keys are presented. A new species is described. WHILE DESCRIBING A new species of Sadieria A red dye for kapa bark cloth was extracted Kaulf., it became obvious to me that there was from the trunk of larger Sadieria species, and confusion regarding circumscription of other occasionally the starchy pith was cooked and species of the genus, their correct names, and eaten (Degener 1930). Palaholo, a rolled-up their descriptions, and that various published and frond of the 'ama 'u fern or a paste made of sap unpublished keys for the genus did not work from the fronds, was used to repair kapa or weld well. In this paper I review and define the five strips of it together (Bryan 1915, Pukui and generally recognized species, describe a sixth, Elbert 1965). and clarify some nomenclatural problems. Sadieria plants are much photographed, fre­ Sadieria is confined to the Hawaiian Islands, quently appearing on magazine covers and tour­ but differs little from Blechnum (Hooker and ist advertisements, and are noted for the pink or Arnott 1832, Hillebrand 1888, Robinson 1913, red young leaves. Bower 1928, Copeland 1947, Joe 1960, Stone 1967, Fosberg 1976). It is usually distinguished from the latter by its pinnate-pinnatifid to bipin­ Taxonomic History nate fronds and its erect to arborescent caudices (Copeland 1947, Stone 1967, Lloyd 1976). How­ Kaulfuss (1824) described the genus Sadieria ever, neither of these characteristics is com­ with the species S. cyatheoides, named in honor pletely lacking in Blechnum, and the arborescent of Dr. Joseph Sadler, a physician who studied caudices are absent in S. squarrosa and S. uni­ the ferns of his native Hungary. Kaulfuss based sora. Robinson (1913) wrote that Sadieria con­ his description on material collected by Adelbert spicuously differed from Blechnum in its greater Chamisso, the naturalist aboard the Russian size, rigidity, and number of scales, but these expedition ship Rurick under the command of are features also shared by some species of Otto von Kotzebue (1821). Since 1824, 11 more Blechnum. Sadieria does differ from other blech­ species and three varieties ofSadieria have been nums in that every species of Sadieria produces described: S. pallida Hook. & Am. (1832) only 16 spores per sporangium. In spite of the [retracted by Hooker (1860), but reinstated by overlapping ofcharacters, all species in Sadieria Hillebrand (1888)]; Blechnum fontanesianum are clearly related to each other and easily recog­ Gaudich. (1827) (=S. cyatheoides); S. (Blech­ nized as a group distinct from Blechnum. num) souleyetiana (Gaudich., l854a) Moore The Hawaiian names (Pukui and Elbert 1965) (1857); S. (Blechnum) squarrosa (Gaudich., for the cyatheoides group are 'ama 'u, ma 'uma 'u, 1854b) Moore (1857); Blechnum kaulfussianum and pua'a 'ehu'ehu ("red pig"), or plural Gaudich. (1854c) (=S. cyatheoides); S. (Blech­ 'ama 'uma 'u. In Hawaiian theology 'ama 'u is num) polystichoides (Brack., 1854) Heller one of the forms that the pig demigod Kama­ (1897) (=S. squarrosa); S. (Polypodium) uni­ pua'a takes at will. The smaller Sadieria squar­ sora (Baker, 1874) Robinson (1913); Gymno­ rosa is called 'apu 'u or 'apulu. gramme sadierioides Underw. (1897) (=S. unisora); S. hillebrandii Robinson (1913) (=S. pallida); S.fauriei Copel. (1914) (=S. pallida); 1 Manuscript accepted 23 October 1996. 2 Herbarium Pacificum, Bishop Museum, Honolulu, S. rigida Copel. (1916) (=S. pallida); S. (Blech­ Hawai'i 96817-0916. num) squarrosa (Gaudich., l854c) Moore 288 A Revision of Sadieria-PALMER 289 (1857) 13 var. tripinnatiJida Hillebr. (=S. squar­ measurements for S. pallida were changed when rosa); S. squarrosa 'Y var. intermedia Hillebr. the two sets ofdata were combined. All measure­ (=S. squarrosa); and S. (Blechnum) squarrosa ments are given as length followed by width. (Gaudich., 1854c) Moore (1857) 3 var. depaup­ erata Hillebr. (=S. unisora). Hillebrand (1888) was the first to publish a Chromosome Number review of the genus, recognizing four species The chromosome number n = 33 (obtained (Sadleria cyatheoides, S. pallida, S. souleyeti­ from Sadleria squarrosa) (Walker 1973, F. S. ana, and S. squarrosa) with three varieties. Rob­ Wagner 1995) falls within the aneuploid series inson (1913) recognized Baker's Polypodium ofBlechnum. The precise number has been diffi­ unisora by adding Sadleria unisora as a fifth cult to obtain because the chromosomes are quite species. She also treated S. pallida Hook. & variable in size, and although they pair nicely, Am. as a synonym for S. cyatheoides Kaulf. and some very large chromosome pairs are confusing described S. hillebrandii as a nomen novum for (F. S. Wagner 1995). S. pallida, thus creating a persisting problem regarding application of these names and confu­ sion about the correct identity of the species Ecology and Distribution involved (Skottsberg 1942, Holbrook-Walker and Lloyd 1973, Degener 1974, Fosberg 1976, Sadleria species are found in habitats ranging Lloyd 1976). from exposed, bare, recent lava flows; to mesic Christensen (1925) recognized seven species, and wet rain forests; and to dark, damp, steep including S. fauriei and S. rigida. Copeland valley walls. Sadleria cyatheoides is among the (1947) and Stone (1967) mentioned that six or first invaders of exposed, new lava flows but seven species existed. In recent years the five is occasionally found in closed-canopy forests. species recognized by Robinson have generally Sadleria pallida is most common in wet, closed­ been accepted by local naturalists and are listed canopy forests but is also found in exposed areas in unpublished fem lists by C. Lamoureux and with high rainfall. Sadleria souleyetiana is most W. H. Wagner, Jr. common in mesic to wet forests but may be found in other habitats. Sadleria squarrosa and S. unisora, as well as S. wagneriana, sp. nov., are limited to dark, moist environments, usually MATERIALS AND METHODS on nearly vertical walls of streams. This revision is based on fieldwork on all the major Hawaiian Islands and study of herbarium Origin and Relationships specimens of MICH and BISH as well as holotypes and other important specimens from K, B, P, GL, Previous authors (Hooker and Arnott 1832, LE, and MPPD. Hillebrand 1888, Robinson 1913, Bower 1928, Dorothy Minard-Elsaesser's thesis (1982) on Copeland 1947, Stone 1967, Fosberg 1976) the systematics ofSadleria was used extensively. accepted the probable monophyletic origin of She examined and measured 1054 herbarium Sadleria from Blechnum as the result of a single specimens and photographs from B, BISH, BM, introduction into the Hawaiian Islands. After the FI, GH, GL, K, MICH, NY, P, UC, and us. Her original introduction the species probably arose measurements were used, with permission, but autochthonously, with geographic isolation and with the following modifications. If my exami­ genetic drift in diverse habitats on the Islands. nation of plants in the field, or from more recent Sadleria can be divided into two groups based collections, differed from hers, I modified the on several characters. (1) The cyatheoides group measurements accordingly. She recognized S. includes S. cyatheoides, S. pallida, S. souleyeti­ rigida as a species distinct from S. pallida, but ana, and S. wagneriana, sp. nov. These species stated that the two might be one species. I recog­ are variable in size but with fronds usually at nize them as one species, and because S. rigida least 0.5 m long, often much longer, and deeply is generally smaller than S. pallida, many of the pinnate-pinnatifid to bipinnate at the bases of 290 PACIFIC SCIENCE, Volume 51, July 1997 larger pinnae. The bases of the sinuses have a such disturbed sites as near road construction narrow, cartilaginous, translucent bar perpendic­ and along trailsides. In some areas intermediates ular to the costa connecting the segments, usu­ are common, with or without disturbance, but ally making them less than completely bipinnate. in other areas they are rare. All except S. wagneriana, sp. nov. lack glands. The spores, without perispore, are smaller than Taxonomic Problems those of the next group: 42.0-70.0 (mean = 53.5) J.Lm long, 26-45 (mean = 35.4) J.Lm wide The taxonomic level at which intraspecific (Lloyd 1976); 50-64 (mean = ca. 55) J.Lm long, diversity in Sadleria is recognized in this paper 31-42 (mean = 34) J.Lm wide (Selling 1946). needs explanation. Hawaiian seed plant species (2) The squarrosa group includes S. squarrosa are noted for their intraspecific variation (Carl­ and S. unisora. Fronds are generally less than quist 1965, 1980, Gillett 1972, W. L. Wagner et 0.5 m long, are fully bipinnate, have glandular aI. 1990). In Hawai'i the endemic ferns also trichomes on the pinnule margins and on the show substantial infraspecific variability, the scales (most easily seen in the field, obscure nonendemic indigenous species show little, and on dried material), and larger spores: 58.0-81.0 recently introduced ferns almost none. Among (mean = 67.5) J.Lm long, 37-45 (mean = 35.4) endemic species there may be marked variation J.Lm wide (Lloyd 1976); 60-75 (mean ca. 67) J.Lm in one or more characteristics (Hillebrand 1888, long, 37-51 (mean = ca. 44) J.Lm wide (Selling Christensen 1925), including frond size; density, 1946). Skottsberg's (1942) measurements of distribution, size, and quality ofscales and hairs; only four spores in this group showed little dif­ degree of blade dissection; sorus size; segment ference in size from his measurements of the size and shape; and depth and shape of sinuses. cyatheoides group. Infraspecific variants are often connected by a cline of intermediate forms. The causes of this variation are not understood, and the variation Hybrids may reflect incomplete speciation or hybridiza­ Plants intermediate between any species pair tion and introgression between species.
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