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Vegetative Anatomy of Dubautia, Argyroxiphiun1j And

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Vegetative Anatomy of Dubautia, Argyroxiphiun1j And Vegetative Anatomy of Dubautia, Argyroxiphiun1J and If/ ilkesia (Compositae) 1 SHERWIN CARLQUIST2 BECAUS E Dubautia, Argyroxipbiam, and lVil­ 442) between Railliardella, a genus tradi­ kesia are endemic Hawaiian genera of uncer­ tionally placed in Senecioneae, and the Juan tain po sition within the Composit ae and are Fernandez Senecioneae Robinsonia and Rbetino­ characterized by species markedly different in dendron. Alth ough the systematic po sitions of habit, a more thorough knowledge of ana­ Argyroxiphimn and lVi/kesia have been in tomical structure in these genera and in doubt, they have been interpreted as belong­ putatively related genera is desirable. The pur­ ing to the tarweeds (Heliantheae, subrribe pose of this study is to explore the variation Madinae) by such authors as Hoffmann (1890: pattern of anatomical characters in vegetative 248). Hoffmann, however, places Dubautia organs of Dubauti«, Argyroxipbium, and lVil­ and Railliardia in the subtribe immediately kesia, and to suggest which of these appear to preceding Madinae, Galinsoginae. Skottsberg be important in indicating rel ationships (1931: 56; 1956: 211) finds Dubautia and among the genera and to other genera. The Railliardia possibly related to Robinsonia and data may also be helpful in outlining natural Rhetinodendron, as well as to a New Guinea groups within the genus Dubautia. genus of Senecioneae, Bracbionostylam. Kec k In formation concerning secondary xylem (1936: 8) agrees, alth ough he emphasizes the of Dubautia is included in a separate study relation of Dubautia to A rgyroxiphiu11l and (Carlquist, 1958). The peculiar leaves of lVilkesia, which he excludes from M adinae Argyroxipbium, and comparison of them with and places in Galinsoginae; and he suggests leaves of lVi/kesia, form the subj ect of an that Dubauti« (sensu lato), A rgyroxipbium, and earlier paper (Carlquist, 1957d ). The present lVilkesia form a related endemic group. Th e paper concerns the vegetative anatomy of the fact that he relates Dubautia to elements of three genera in other respects. For the pur­ both Heli antheae and Senecioneae is not in­ poses of this study, the writer interprets Du­ con gruent, in his opinion, because Dubauti« bautia as including Railliardia, a treatment may be regarded as a genus transitional be­ offered by Keck (1936: 24- 28) and accepted tween the tribes . Dubautia, therefore, would by St. John (1950). seem to be a critical genus in the delimitation The dubious position of Dubauti« can be of the tribes of Compositae. St. John (1950: appreciated by comparing the works of au ­ 240) finds no genus with close affinities to thors who have dealt with the genus. Al­ Dubautia, and Sherff (1935) declines to com­ though recent authors agree that Dubautia ment. Because the Heliantheae mentioned and Railliardia, if both genera are recognized, above are American, demonstration of rela­ are closely related, earlier authors recognized tionship between them and Dubautia, Argyro­ both genera and relegated them to widely xipbium, and lVi/kesia would be important by separated portions of the family. Bentham validating an example of Hawaiian-American and H ooker (1873: 393), for example, place phytogeo graphical affinity. Dubautia near A rgyroxiphiu11l and lVi/kesia, MATERIALSAND METHODS whereas they locate Railliardia (loc. cit., p. The author's collections (listed below) I Manuscript received November 20, 1957. from which portions were preserved in Car­ 2 Claremont Graduate School, Rancho Santa Ana Boranic Garden, Claremo nt, Californ ia. noy 's Fluid (3 parts absolute eth yl alcoho l: 195 196 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 H 20a (UC); D. platyphylla (Gray) Keck, species, however, were studied from frag­ Forbes 1101M (DC); D. railliardioides Hillebr., ments of herbarium specimens treated accord­ Carlquist H16 (DC); D. reticulata (Sherff) ing to techniques described earlier (Carlquist, Keck, Rock 8573 (DC); 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 H 20 (DC); D.Sherfftana Fosb ., scribed in that paper. Preparations made from St.John 23924 (BISH);D.strutbioloides (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); D. waialealae Rock, Rock Oct. 1911 analyze structure of several portions of a leaf, (GH, cotype). Argyroxiphimn Caliginii Forbes, and to secure stems both in primary condi­ Carlquist H 28 (DC); A . Grayanum (Hillebr.) tion, for a study of stem structure, and with Degener, Carlquist H27 (DC); A . sand­ some secondary growth, for stud y of mature wichense DC, Carlquist H19 (DC) ; Wilkesia pith types. Pith of the rosette species of gymnoxiphium Gray, Carlquist HlO (DC). Argyroxipbium (A . sandwichense) was taken Gratitu de is expressed to the curators of from basal (epicotyl) portions of the rosette. these herbaria for use of their materials. Data Leaf anatomy was studied in all species except concerning the 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 Dubautia and A rgyroxiphium, as in the Hawaiian Islands. well as for Wilkes/a. Subspecific variation was ANAT OMY disregarded in this study, and analysis of the LeafTransections hybrids recognized by Sherff (1935) did not seem feasible at present . Characters of leaf anatomy as seen in rran­ Dubautia arborea (Gray) Keck, Rock 8344 sections constitute the best indications of (UC); D. ciliolata (DC) Keck, var. laxiflora specific diversity in Dubautia. These charac­ (DC) Keck, Rock 10326 (UC); D. Hille­ ters include bifacial or isolateral organization, brandii (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. laevigata Gray, Heller width of palisade, frequency of bundle­ 2616 (UC); D. latifolia (Gray) Keck, Heller sheath extensions, presence of fibers within 2887 (UC); D. laxa H . & A:, Carlquist H1 2 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. J ohn uniseriate non glandular or biseriate glandular 10303 (UC) ; D. magnifolia Sherff, Rock 9012 hairs. Cuticle is not included in description of (BISH); D. Menziesii (Gray) Keck , Carlquist epidermis cell wall thickness, because it was H17 (UC); D. microcephala Skottsb., Carl­ invariably a very thin layer. Numerous prep­ quist H14 (UC); D. molokaiensis (Hillebr.) arations showed separation of the cuticle from Keck, Forbes 86Mo (UC); D. montana (H . the epidermis (e.g.,Fig. 3). The species of Mann) Keck, Rock 8594 (UC); D. paleata Dubautia are compared below by reference to Gray, Forbes 914K (BISH) ; D. plantaginea " types" based on the species illustrated. No Vegetative Anatomy - CARLQUIST 197 taxonomic sig nificance is necessaril y implied 1. D. Knudsenii (Fig . 1). Leafbifacial; cells of by this arrangement. The gro ups below, how­ upper epidermis larger than those oflower 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 m ajor veins only; fibro us bundle cap species most closely resembling it are listed present only on major veins ; secretory beneath, with the features by which the y canals 4 near larger veins, 1 or 2 near smal­ differ mentioned in parentheses. ler veins; trichomes absent at maturity. FIGS. 1- 6. Leaves of Dubautia. (1-5) Leaf transecrions . (1) D . Knudsenii, X 75. (2) D. railliardioides, X 67. (3) D. latifolia , X 95. (4) D. waialealae, X 87. (5) D . M enziesii, X 67. (6) D. loncbopbylla, po rtion of cleared leaf to show marginal trichomes, below, and secreto ry canals, which appear as darker lines (on account of resin s) beside two larger veins, X 58. 198 PACIFIC SCIENCE, Vol. XIII , April, 1959 Like D. Knudsenii: 3. D.latifolia (Fig . 3). Leafbifacial, 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. reticulate (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 fibers 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;
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