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Rhe Peculiar Leaves Omparison of Them with Form the Subject of an Ist. 1957

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Rhe Peculiar Leaves Omparison of Them with Form the Subject of an Ist. 1957 ... „ —J ... -J rhe peculiar leaves of Wilkesia, which he excludes from Madinae omparison of them with and places in Galinsoginae; and he suggests form the subject of an that Dt/baiitia (sens// lato), Argyroxiphium, and ist. 1957^). The present Wilkesia form a related endemic group. The egetative anatomy of the fact that he relates Di/buutia to elements of n respects. For the pur- both Heliantheae and Senecioneae is not in- the writer interprets Du- congruent, in his opinion, because Dubautia 196 PACIFIC SCIENCE, Vol. XIII, April, 1959 1 part glacial acetic acid) provided good ma­ Gaud. var. platyphylla Hillebr.. Carlquist terial for anatomical study. The others of the H20a (UC); D. platyphylla (Gray) Keck, species, however, were studied from frag­ Forbes 1101M (UC); D. railliardioidesHillebr., ments of herbarium specimens treated accord­ Carlquist Hl6 (UC); D. reticulata (Sherff) ing to techniques described earlier (Carlquist, Keck, Rock 8573 (UC); D. Rockii (Sherff) 1957a: 207). The liquid-preserved material Keck, Rock 8601 (BISH); D. scabra (DC) was prepared by means of techniques de­ Keck, Carlquist H20 (UC); D. Sherffiana Fosb., scribed in that paper. Preparations made from St.John 23924 (BISH); D.struthioloides (Gray) herbarium material, both as whole mounts Keck, Wilkes Exped. s. n. (GH, type); D. and as sections, gave quite satisfactory infor­ ternifolia (Sherff) Keck, Forbes 1175 (BISH); mation concerning vegetative anatomy. Every D. thyrsiflora (Sherff) Keck, Forbes 1203M effort was made to secure mature leaves, to (GH : /). waialealae Rock, Rock Oct. L$>11 analyze structure of several portions of a leaf, (GH, cotype). Argyroxiphinm Caliginii Forbes, and to secure stems both in primary condi­ Carlquist H28 (UC); A. Grayanum (Hillebr.) tion, for a study of stem structure, and with Degener, Carlquist H27 (UC); A. sand­ some secondary growth, for study ot mature wichense DC, Carlquist H19 (UC); Wilkesia pith types. Pith of the rosette species of gymnoxiphium Gray, Carlquist H10 (UC). Ariyroriphium {A. sandwichense) was taken Gratitude is expressed to the curators of from basal (epicotyl) portions ot the rosette. these herbaria for use of their materials. Data Leat anatomy was studied in all species except concerning die stem of D. platyphylla were D. coriacea (Sherff) Keck, D. demissifolia obtained from a slide (prepared from a liquid- (Sherff) Keck, and D. kohalae (Skottsb.) St. preserved collection by Degener, no. 19188) John. These species would probably add little kindly given to the writer by Dr. John W. to the gamut of variation presented here. Hall. Thanks are due Dr. Harold St. John for Stem anatomy was studied for the majority of his assistance during the author"s field work the species of Duhautia and Argyroxiphium, as in the Hawaiian Islands. well as tor Wilkesia. Subspecific variation was ANATOMY disregarded in this study, and analysis of the hybrids recognized by Sherff (1935.) did not sections seem feasible at present. Characters of leaf anatomy as seen in tran­ Duhautia arborea (Gray) Keck, Rock 8344 sections constitute the best indications of (UC); D. ciliolata (DC.) Keck, var. laxiflora specific diversity in Duhautia. These charac­ (DC) Keck, Rock 10326 (UC); D. Hille- ters include bifacial or isolateral organization, brandit (H. Mann) Keck, Hillebrand s.n. relative size of cells in upper and lower epi­ (GH, syntype); D. Knudsenii Hillebr., Carl­ dermis, thickness of cell walls in epidermis, quist H15 (UC); D. Gray, Heller width of palisade, frequency ot bundle- 2616 (UC); D. latifolia (Gray) Keck, Heller sheath extensions, presence of fibers within 2887 (UC); D. laxa H. & A., Carlquist H12 the bundle sheath, presence of secretory ca­ (UC); D. linearis (Gaud.) Keck, Rock 8123 nals in the bundle sheath, and occurrence of (UC); D. lonchophylla (Sherff) Keck, St. John uniseriate nonglandular or biseriate glandular 10303 (UC); D. magnifolia Sherff, Rock 9012 hairs. Cuticle is not included in description ot (BISH); D. Menziesii (Gray) Keck, Carlquist epidermis cell wall thickness, because it was invariably a very thin layer. Numerous prep­ H17 (UC); D. microcephala Skottsb., Carl­ arations showed separation ot the cuticle from quist H14 (UC); D. molokaknsis (Hillebr.: the epidermis (e.g., Fig. 3). The species of Keck, Forbes 86Mo (UC); D. montana (H. Duhautia are compared below by reference to Mann) Keck, Rock 8594 (UC); D. paleata "types" based on the species illustrated. No Gray, Forbes 914K (BISH); D. plantaginea Vegetative Anatomy — CARLQL'IST 197 taxonomic significance is necessarily implied D. Knudsenii (Fig. 1). Leaf bifacial; cells of by this arrangement. The groups below, how­ upper epidermis larger than those of lower ever, do appear to be natural groups in some epidermis, thin walled; 1 layer of palisade instances. For each group, the illustrated spe­ present; bundle-sheath extensions present cies is used as a basis for description, and the on major veins only; fibrous bundle cap species most closely resembling it are listed present only on major veins; secretory- beneath, with the features by which they canals 4 near larger veins, 1 or 2 near smal­ differ mentioned in parentheses. ler veins; trichomes absent at maturity. FlGS. 1-6. Leaves of Dubatitia. (1-5) Leaf transections. (1) D. Knudsenii. X 75. (2) D. railliardioides. X 67. (3) D. latifolia, X 95. (4) D. uaialealae, X 87. (5) D. Menziesii, X 67. (6) D. lomhophylla, portion of cleared leaf to show marginal trichomes. below, and secretory canals, which appear as darker lines (on account of resins) beside two larger veins, X 58. 198 PACIFIC SCIENCE, Vol. XIII, April, 1959 Like D. Knudsenii: D. latifolia (Fig. 3). Leaf bifacial, thin; up­ D. laxa (a few uniseriate hairs on lower per and lower epidermis cells of approxi­ surface). mately the same size, very thick walled: a D. molokaiensis (thicker outer walls on single layer of cells weakly defined as upper epidermis). palisade; bundle-sheath extensions fre­ D. reticulata (thick-walled upper epi­ quent, consisting of isodiametric to fiber­ dermis; uniseriate hairs frequent on like sclereids, although no "bundle cap" both surfaces, especially in the libers are present on the vascular bundles: grooves overlying veins on the 1 or 2 secretory canals present within the upper surface). bundle sheath or sheath extension of many- 2. D. railliardioides (Fig. 2). Leaf bifacial; veins; hairs absent at maturity. cells ot upper epidermis larger than those Like D. latifolia: of lower epidermis; prominent grooves on D. laevigata (sclerenchyma of bundle upper surface above major veins; 2 pali­ sheaths and sheath extensions con­ sade layers present; sheath extensions sisting of thick-walled fibers; secre­ present on many larger veins; fibers com­ tory canals absent'. mon at both poles of larger veins; 4 secre­ i. D. uaialealae (Fig. 4). Leaf bifacial; cells tory canals in outer portion of bundle of upper and lower epidermis of about the sheath of larger veins, 1 or 2 near smaller same size, very thick walled; 1 or 2 layers veins; scattered uniseriate trichomes on somewhat differentiated as palisade; sheath lower surface and margins (upwardly extensions present only on midvein; prom- appressed). nent strands of thick-walled fibers on Like D. railliardioides: phloem face of veins, also on xylem pole D. magnifolia, D. microcephala, D. of larger veins; secretory canals absent; plantaginea (virtually identical with large uniseriate trichomes abundant on D. railliardioides). upper surface and on margins. D. paleata (sheath extensions on ma­ D. Menziesii (Fig. 5). Leaf somewhat iso- jor veins only: fibers more common lateral (stomata frequent on upper surface; at phloem pole of veins; secretory- adaxial palisade cells short, nearly isodia­ canals absent; numerous uniseriate metric; cells on abaxial face like adaxial hairs on both surfaces). palisade cells in shape, with more nu­ D. ternifolia (fibers more common on merous chloroplasts than cells in the cen­ phloem face of bundles). tral portion of the leaf); leaf very thick; D. scabra, D. ciliolata, D. lonchophylla, bundle-sheath extension only on midvein; D. linearis, D. Sherjfiana, D. thyrsi- large groups of fibers on phloem face of flora (upper epidermis very large veins (a few libers on xylem pole of celled; bundle-sheath extension on larger veins); secretory canals infrequent, midvein only; fibers mostly on 1 or 2 in bundle sheath ot larger veins phloem face of bundle; usually 1 near leaf base; thick-walled uniseriate secretory canal above xylem pole of trichomes scattered over both surfaces and larger veins). margin of leaf. D. platyphylla (fibers at phloem face of Like D. Menziesii: bundles only; secretory canals ab­ D. Hillebrandii (nearly identical with sent; biseriate glandular hairs with D. Menziesii). large multicellular capitate heads D. arborea (fibers lacking near veins; and long stalks abundant on both biseriate glandular hairs present on surfaces; uniseriate hairs absent). both leaf surfaces). Vegetative Anatomy — CARLQUIST 199 D. Rockii (a single secretory canal impracticable because the acid released such adaxial to most veins). compounds from the wall and caused swelling D. montana (fiber strands near veins and distortion of the leaves. The phenomenon small :. described by the writer as "abaxial meristem" D. struthioloides (more nearly isolateral in Argyroxiphium is totally lacking in Du­ than D. Menziesii). bautia, as it is in Wilkesia. The different types of leaf structure de­ Leaf Venation tailed above reflect ecological conditions for Dubautia to a certain extent, as well as being Figures 7-9 show the range of venation indicative of taxonomic groupings. The thin, types which may be seen in the genus Du­ glabrous leaves of D. Knudsenii, with thin- bautia. Dubautia latifolia (Fig. 7) is excep­ walled epidermal cells, correspond to the tional for the genus in its reticulate vein pat­ moist forest areas on Kauai where that spe­ tern with polygonal areoles containing nu­ cies grows. On the contrary, the tendency merous freely-terminating veins. Except for the toward thick, isolateral leaves with thick- major veins (two of which are shown, left and walled epidermal cells in D.
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