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INTERVASCULAR PIT MEMBRANE STRUCTURE in DAPHNE and WIKSTROEMIA - SYSTEMATIC IMPLICATIONS 1 by Roland R

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INTERVASCULAR PIT MEMBRANE STRUCTURE in DAPHNE and WIKSTROEMIA - SYSTEMATIC IMPLICATIONS 1 by Roland R IAWA Journal, Vol. 17 (2),1996: 161-181 INTERVASCULAR PIT MEMBRANE STRUCTURE IN DAPHNE AND WIKSTROEMIA - SYSTEMATIC IMPLICATIONS 1 by Roland R. Dute-, John D. Freeman", Frank Henning- & Logan D. Barnard- SUMMARY Intervascular pit membranes were investigated in species of Daphne, Wikstroemia, and other allied genera of the Thymelaeaceae. Results con­ firmed a previous study showing that, except for section Mezereum, all sections of Daphne had pit membranes with tori. Taxonomically isolated species D. aurantiaca and D. genkwa had tori, but lacked a G-layer. Tori similar in structure to those of D. aurantiaca and D. genkwa were ob­ served in three species from the subgenus Diplomorpha of Wikstroemia. Tori of a slightly different morphology were noted in W kudoi (subg. Daphnimorpha). Tori appeared absent from species of the subgenus Wikstroemia (= Euwikstroemia of Domke), and from the genera Drapetes, Edgeworthia, and Eriosolena. These results suggest a close relationship between Daphne and Wikstroemia. The degree of torus development and the distinctiveness of helical thickenings suggest that smaller tracheary elements serve as a backup water-conducting system to larger vessel elements. Key words: Daphne, Wikstroemia, Thymelaeaceae, pit membrane, torus, wood ultrastructure. INTRODUCTION The genus Daphne is unusual in that it is one of four genera ofangiosperms known to have intervascular pit membranes with tori (q.v. discussion in Dute et al. 1990). Ohtani and Ishida (1978a) were the first to comment upon this feature in three species of Daphne common to Japan. Later, Dute et al. (1990) detailed torus development in D. odora . In a survey of five species, Ohtani (1983) observed that the three species with tori belonged to a different section of the genus than the two species without tori. Dute et al. (1990) independently noted that whereas intervascular pit membranes in D. odora and D. cneorum always had tori, those of D. mezereum did not. It was hypothesized 1) Contribution No. 6-955054 from the Alabama Agricultural Experiment Station, Auburn University, Auburn, AL 36849, U,S. A. 2) Department of Botany and Microbiology and Alabama Agricultural Experiment Station, Auburn University, Auburn AL 36849, U.S .A. Correspondence should be addressed to R.R. Dute. 3) College of Agricultural and Environmental Sciences, Cooperative Extension Service, The University of Georgia. 4) Department of Horticulture, Auburn University. Downloaded from Brill.com10/06/2021 02:44:15PM via free access 162 IAWA Journal, Vol. 17 (2), 1996 that the presence of torus-bearing pit membranes could be correlated with either growth habit (D. odora and D. cneorum are evergreen; D. mezereum, deciduous) or systemat­ ics. A later study (Dute et al. 1992) of 22 taxa (21 species and 1 hybrid) of Daphne seemed to confirm the latter possibility. The only species without tori were those with­ in the section Mezereum. Nevertheless, because Daphne has 50 to 70 species (Bailey 1949; Brickell & Mathew 1976) the study of Dute et al. (1992) cannot be considered comprehensive. The species so far investigated are listed in Table 1. Table I. Distribution of common horticultural species within the genus Daphne according to Brickell and Mathew (1976) based on Kessler (1898) . Species with name s in boldface type previously were examined for presence of tori in this laboratory (Duteet al. 1990, 1992). Daphne (Daphne Sect. Mezereum sect. Daphnanthes subsect. Oleoides contd) D.jezoensis D. rodriguezii D. kamtschatica D. stapfi i D. koreana D. transcaucasica D. mezereum Subsect. Gnidium D. pseudomezereum D.gnidium D. rechingeri Subsect. Collinae Sect. Genkwa D. blagayana D. genkwa D. collina Sect. Laureola D. macrantha D. albowiana D. sericea D. glomerata Subsec t. Cneorum D.laureola D. arbuscula D. pontica D. cneorum Sect. Eriosolena D.juliae D. aurantiaca D. petraea Sect. Daphnanthes D. striata Subsect. Alpinae Subsect. Daphnanthoides D. alpina D. acutiloba D. altaica D. bholua D. caucasica D. grueningiana D. giraldii D. kiusiana D. sophia D. longilobata Subsect. Oleoides D.luzonica D. baksanica D. miyabeana D. euboica D.odora D. gnidioides D. papyracea D.jasminea D. retusa D. kosaninii D. shillong D. linearifolia D. sureil D. mucronata D. taiwaniana D.oleoides D. tangutica Downloaded from Brill.com10/06/2021 02:44:15PM via free access Dute, Freeman , Henning & Barnard - Pit membranes in Daphne and Wikstroemia 163 The genus Daphne is notwell circumscribed.Species of section Eriosolena are some­ times included in a separate genus due to their distinct floral characteristics (e.g., Hallier 1922; Leandri 1930; Domke 1934). Other species, mostly of the section Genkwa, have been placed in the genus Wikstroemia, based in part upon opposite leaf arrangement. The purpose of this study is to continue the survey of Daphne species for torus­ containing pit membranes. Some species of Wikstroemia and other related genera (Dra­ petes, Edgeworthia, Eriosolena) also are examined. Although no one feature provides sufficient grounds on which to base a taxonomic treatment, pit membrane structure in conjunction with floral and vegetative characters furnishes valuable information for understanding the circumscription of Daphne. MATERIALS AND METHODS Table 2 lists the species used in this study. Herbarium specimens were provided by Dr. Hiroo Kanai of the Tokyo National Science Museum, Dr. Mikio Ono of the Makino Herbarium of the Tokyo Metropolitan University, Drs. Ghillean T. Prance and G. L1. Lucas of the Royal Botanic Gardens, Kew, and Dr. P. Baas of the Rijksherbarium, Leiden . A single specimen of Daphne genkwa from the Auburn University Herbarium was also studied. Dr. Stephanie Mayer of the University of Chicago supplied air-dried branches from living specimens in her collection. These collections provided exam­ ples of species from the closely related genera Daphne, Wikstroemia , Drapetes, and Eriosolena . In addition, living specimens of another genus of the Thymelaeaceae, Edge­ worthia, were obtained from Gossler Farms Nursery, Springfield, Oregon, USA and grown in a greenhouse until needed for comparative purposes. For scanning electron microscopy (SEM), split radial sections (from branches 3-5 mm in diameter) of herbarium specimens were attached to aluminum stubs with dou­ ble stick carbon tape. Specimens and stubs were sputter-coated with gold-palladium. Split radial sections ofEdgeworthia papyrijera were allowed to air-dry before mount­ ing. Specimens were viewed with a Zeiss Digital Scanning Microscope (DSM 940) at voltages of 5, 10, and 15 kY. Preparation of herbarium specimens for transmission electron microscopy (TEM) was according to the method of Dute et al. (1992) in which pieces of wood no more than I mm on a side were soaked in three changes of absolute acetone followed by propylene oxide and embedment in Spurr's resin (Spurr 1969). Transverse, radial and tangential monitor sections of 1.5-2 mm were cut, affixed to glass slides, and stained with toluidine blue 0 and basic fuchsin for light microscopy. Transverse silver sec­ tions were cut on a Sorvall MT-2b ultramicrotome and stained with uranyl acetate and lead citrate . Observations were made with a Zeiss 10 transmission electron micro­ scope operated at 60 kV. Monitor sections often would not dry flat onto a glass slide but rather contained air bubbles . To eliminate this problem and to provide better resolution, monitor sections were deplasticized using the following method .Transverse sections of Spurr's-embed­ ded material were cut to thicknesses of 0.5 to 3 mm and placed into a drop of water on a precleaned circular coverslip (15 mm diameter). The sections then were heat-fixed to Downloaded from Brill.com10/06/2021 02:44:15PM via free access 164 IAWA Journal, Vol. 17 (2),1996 Table 2. Sources of wood specimens examined in this study. Taxon Herb./City Collection date Collector(s) number Daphne Sect. Mezereum D. kamts chaticum TNS 10 Jul1933 Ono s.n. D. koreana TNS 11 Aug 1954 Furuse s.n . Sect. Genkwa D. genkwa Lw < 1827 Siebold & Zuccarini s.n. K 21 Apr 1981 Green 2148 AUA 9 May 1980 Miller & Yinger Sect. Eriosolena D. aurantiaca K 1913 Forrest 1018 D.pendula K 1 Apr 1930 Henderson 23274 Sect. Daphnanthes Subsect. Alpinae D. alpina TNS I Jun 1903 Bouchard s.n. D. altaica TNS 1840-1841 Schrenk 1306 Subsect. Oleoides D. gnidiodes K 26 Jul1960 Khan, Prance & Ratcliffe 255 D.jasminea K Apr 1939 Atchley 1866 D. stapfii K None listed Jacobs 6604 Subsect. Gnidium D. gnidium TNS 6 Oct 1950 Matos & Matos s. n. Subsect. Cneorum D. bholua TNS 9 Mar 1953 Nakao s.n . K 29 May 1971 Colville , Barclay & Syrge 2396 D. petraea K 24 May 1870 Porta s.n. Subsect. Daphnanthoides D. arisanensis TNS 20Aug 1969 Yamazaki, Namba & Tani s.n. D. kiusiana TNS 19 May 1974 Ito 234 D. miyabeana TNS 7-9 Aug 1958 Okuyama 12837 Subsect. Collinae D. collina TNS 26 Jun 1853 Kotschy s.n. Drapetes D. ericoides TNS 24 Jun 1974 Craven 2768 Eriosolena E. wallichii K None listed Hooker 11150 Edgeworthia E. papyrifera Auburn 15 Oct 1992 Dute Wikstroemia Subg . Diplomorpha W. albiflora MAK 16-20 Aug 1960 Hatusima & Sako 25120 W. ganpi MAK Sep 1936 Makino s.n. W. pauciflora MAK Aug 1904 Makino s.n . W. sikokiana MAK 10 Jul1956 Satomi s.n . W. trichotoma MAK 22 Ju11964 Murata 18605 W. yakushimensis MAK 16-20 Aug 1960 Hatusima & Sako 25075 Subg . Daphnimorpha W. kudoi MAK 20Jul1965 Satomi 25126 Subg. Wikstroemia W. pseudoretusa MAK 12 Nov 1970 Ono & Kobayashi s.n. W. retusa MAK 19 Mar 1981 Sugawara 1565 Chicago 21 Sep 1994 Mayer W. indica Chicago 21 Sep 1994 Mayer Downloaded from Brill.com10/06/2021 02:44:15PM via free access Dute, Freeman, Hennin g & Barnard - Pit membranes in Daphne and Wikstroemia 165 the coverslip. Resin was removed using a Poly/Solv Kit (Polysciences Inc., Warrington, PA, USA, Data Sheet #291). The instructions were followed except that it was found necessary to increase the time in the initial plastic solubilizing mixture from 5 to 10 minutes.
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