Ken Ogata Forestry & Forest Products Research Institute, P.O
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IAWA Bulletin n.s., Vol. 12 (2),1991: 111-121 WOOD ANATOMY OF ZABELIA (CAPRIFOLIACEAE): EVIDENCE FOR GENERIC RECOGNITION by Ken Ogata Forestry & Forest Products Research Institute, P.O. Box 16, Tsukuba Norin Kenkyu Danchi, Ibaraki, 305 Japan Summary The wood anatomy of Zabelia is described Zabelia (Rehder) Makino ex Hisauchi et Hara based on stem and twig samples. Wood anat Series 1. Zabelia omy supports the recognition of Zabelia 1. Z. angustifolia (Bur. et Franch.) Makino: often considered to be congeneric with Abelia W. China. - as a distinct genus, on the basis of type of 2. Z. buddleioides (W.W. Smith) Hisauchi perforations, vessel grouping and porosity. et Hara: W. China. Zabelia is compared with other woody genera 3. Z. corymbosa (Regel et Schmalh.) Maki of the Caprifoliaceae using selected wood no: Turkestan. anatomical features (Table 2). The presence 4. Z. mosanensis (Chung ex Nakai) Hisa of aggregate rays is a very distinguishing fea uchi et Hara: Korea. ture of Zabelia not found in any other genus 5. Z. triflora (R.Br.) Makino: W. Himalaya. of the Caprifoliaceae. 6. Z. ryaihyonii (Nakai) Hisauchi et Hara: Key words: Caprifoliaceae, Zabelia, wood Korea. anatomy, taxonomy. Series 2. Biflorae (Zabel) Hara 7. Z. biflora (Turcz.) Makino: N. China. Introduction 8. Z. coreana (Nakai) Hisauchi et Hara (= In a previous paper on the wood anatomy Z. biflora var. coreana (Nakai) Hara): N. of Japanese Caprifoliaceae (Ogata 1988), Korea to Ussuri. herbarium twig samples of Zabelia integri 9. Z. die/sii (Graebn.) Makino (= Abelia folia were examined. It was tentatively con anhwensis Nakai, A. brachystemon cluded that the characters of Z. integrifolia (Diels) Rehder, A. hersii Nakai, A. on were closer to those of a group of Lonicera kocarpa (Graebn.) Rehder, A. zanderi species ('Lonicera B') than to those of Abelia. (Graebn.) Rehder): Central to W. China. Zabelia has generally been considered to be 10. Z. insularis (Nakai) Hisauchi et Hara (= close to, or even congeneric with Abelia es Z. biflora var. coreana f. insularis (Na pecially in most of the Chinese and Korean kai) Hara): S. Korea. literature. The wood anatomical data on II. Z. integrifolia (Koidz.) Makino: Japan. Zabelia were, however, insufficient to dis 12. Z. umbellata (Graebn. et Buchw.) Ma cuss the problem at that time, because they kino: Central to W. China. were based on twig samples of only one spe cies. Of these species, I recently could obtain good According to Hara (1983), the genus stem wood samples of four species: Zabelia Zabelia consists of two series and 12 spe biflora, Z. dielsU, Z. integrifolia, and Z. mo cies, which are distributed from Afghanistan, sanensis, and also twig samples of these Turkestan and western Himalaya through and six other species taken from herbarium China east to Korea, southern Ussuri and sheets. Twig samples could be used satisfac Japan: torily for checking porosity, vessel grouping, Downloaded from Brill.com09/26/2021 06:40:36AM via free access .....tv Table 1. Variation in some quantitative wood anatomical characters of Zabelia. Vessels Fibre Ray s Species Frequency %of Tang. diam. ().lIII) Element length ij!m) length (!tm) (TWTwno.) (per mm2) solitary max. width pores earlywood latewood mean (range) ± S.D. mean (range) ± S. D. max. height frequency ---------- (cells) ij!m) (per mm) Zabelia biflora (14147) 350-427 81-93 40-70 20-50 373 (255 -445) ± 45 750 (500- 930) ± 98 4 2900 19-25 dielsii (2778) 201-296 90-96 30-55 15-30 529 (335 -675) ± 83 969 (650-1270) ± 147 3 2100 18-20 dielsii (2780) 259-566 66-82 20-45 15-20 401 (285-525) ± 69 731 (500- 890) ± 93 5 2800 21-25 '-):- integrifolia (14140) 135-288 81-95 25-50 15-30 341 (250-435) ± 46 709 (580- 890) ± 78 4 1600 19-24 I~ 0:; Downloaded fromBrill.com09/26/2021 06:40:36AM mosanensis (2703) 737-741 75-82 20-45 10-20 305 (235 - 750) ± 33 543 (400- 700) ± 66 4 1300 19-25 I§ ('1> g. sp, (2779) 259-328 87-94 40-60 20-40 381 (270-490) ± 64 793 (550- 950) ± 83 3 1200 20-25 I? ~ -< ?- .....tv ~ ..... via freeaccess 'D 'D ..... Ogata - Wood anatomy of Zabelia 113 type of perforation plate, spiral thickenings in al. 834284*. -Z. tyaihyonii: Korea (TWTw vessel and fibres, and aggregate rays, which 2784, diam. 6 mm); Korea, Chungchong Puk are important characters for generic separa Do, T. Nakai 8205*. - Zabelia sp.: China tion of the Caprifoliaceae. (TWTw 2779, 20 mm). The aim of this paper is to present the first 2) Other genera of Caprifoliaceae (all sam complete wood anatomical description of Za ples obtained from herbarium sheets). - Dier belia and also to demonstrate that Zabelia is a villa cf. lonicera Mill.: USA, Massachusetts, genus distinct from Abelia or any other genus T. Shimizu et aI. 35073. - D. rivularis Gatt.: of the Caprifoliaceae. Japan, Tokyo (cult.). - Dipelta floribunda Maxim.: China, Hupeh, A. Henry 6558. -D. Materials and Methods yunnanensis Franch.: SW. China, G. Forrest Microscopic slides were made of stem 5746; China, Yunnan, B. Bartholomew et aI. wood samples according to standard tech 1208.-Heptacodiumjasminoides Airy Shaw: niques, and stained with safranin and fast Japan, Tokyo (cult.), Y. Ito s.n. - Kolk green. Free hand-sectioned, glycerin-mount witzia amabilis Graebn.: Japan, Osaka (cult.), ed temporary slides were made of twig sam G. Murata et aI. 39. - Leycesteria formosa ples from herbarium sheets deposited at the Wall.: India, Darjeeling, H. Kanai et al. University Museum, University of Tokyo. 6253; Japan, Kyoto (cult.), H. Hara s.n. - Macerations of all samples were prepared L. glaucophylla (Hook. f. et Th.) Hook. f.: according to Jeffrey's method. Methods for E. Nepal, H. Hara et al. 6302968. - L. gra measurement of the number of vessels and cilis (Kurz) Airy Shaw: Darjeeling, H. Kanai the length of vessel members and fibres et al. 6254. - L. stipulata (Hook. f. et Th.) are those given in my previous paper. The Fritsch: Darjeeling, H. Hara et al. 6258. - IAWA List (IAWA Committee 1989) was Linnaea borealis L.: Japan, Mt. Yatsugatake, mostly followed for selection and description T. Inokuma s.n.; Japan, Hokkaido, T. Ino of features. kuma s. n. (2 samples). - Symphoricarpos Wood samples examined are as follows: albus Blake: Hokkaido (cult.); Germany, 1) Zabelia (samples marked with * are Hannover (cult.), K. Emura 1198. - S. mi those of twigs from herbarium sheets). - Z. crophyllus Kunth: Mexico, Hidalgo, C.G. biflora: China, Beijing, H. Chu s. n. (TWTw Pringle 13010. - S. orbiculatus Moench: 14147, diam. 28 mm); China, Shanxi, M. USA, Missouri, P. H. Raven et al. 26902. - Togashi s.n.*; Beijing, K.S. Chow 82015* S. utahensis Rydb.: USA, Utah, L. C. An & T. Nakai et aI. s.n.* (2 samples). - Z. derson 1389. buddleioides: China, Yangtze Bend, G. For rest s. n. *. - Z. coreana: Korea (TWTw Generic description 2782, diam. 6 mm); Korea, Kangwon Do, Macroscopic features - Bark generally T. Nakai 13969* (2 samples); Korea, Mt. greyish brown, 0.5-2 mm thick, not papery, Wha-Ak, Y.N. Lee & Y.S. Lee 231*. -Z. with six distinct longitudinal furrows in corymbosa: Kirghizia*. - Z. dielsii: China branches, which often become indistinct in (TWTw 2778, diam. 21 mm); China (TWTw old stems because of the fissured or cracked 2780, diam. 20 mm); China, Shensi, W. bark. In cross section of stems and branches, Purdom 434*. - Z. insularis: Korea, Ullun the furrows can be traced to the central pith as Do lsI., T. Nakai 4578* & 4579* (2 sam radial broad lines (Figs. 1-9), representing ples). - Z. integrifolia: Japan, Mt. Buko, K. aggregate rays. Pith solid, white. Sapwood Ogata s.n. (TWTw 14140, diam. 36 mm); and heartwood not differentiated, pale Japan, Kawakamimura, S. Kurata s.n. *; yellowish grey to pale yellowish brown. Japan, Mt. Ishidate, S. Kurata s. n. *; Japan, Texture fine. Growth rings visible to the Yamaemura, M. Otomasu s.n.*. -Z. mosa naked eye or with a lens. Air dry specific nensis: Korea (TWTw 2783, diam. 17 mm); gravity 0.74-0.95. Korea,Pyongannam Do, T.H.Chong s.n.*. Microscopic features (see also Table 1) - Z. triflora: India, Himachal Pradesh, N. C. Growth rings generally distinct, marked by Nair s.n."'; Nepal, Kali Gandaki, H. Ohba et radially compressed latewood fibres and a Downloaded from Brill.com09/26/2021 06:40:36AM via free access 114 IAWA Bulletin n.s., Vol. 12 (2),1991 Figs. 1-9. CS of stems or twigs showing six regular furrows and broad lines from the centre to periphery. - I & 2: Zabelia biflora (1: stem, x I; 2: twig, xc. 8). - 3: Z. coreana (twig, x c. 8).- 4: Z. corymbosa (twig, x c. 8). - 5: Z. insularis (twig, x c. 8). - 6: Z. integrifolia (stem, x c. 2). - 7: Z. mosanensis (twig, xc. 8). - 8: Z. triflora (twig, x c. 8). - 9: Z. tyaihyonii (twig, x c. 8). few rows of larger earlywood vessels. Wood erally high (Table I); pores round or oval to ring-porous to semi-ring-porous with a few angular; walls 1.5-2 (1-2.5) )lm thick. Per (usually 1-2) rows of larger pores in the forations mostly simple, rarely (0.3-5%) early wood which gradually become smaller scalariform with very few (usually 1, some through the intennediate to the latewood times 2) bars; bars often incomplete or ves (Figs. 10, II, 14, 18 & 19), sometimes near tigial (Fig. 23); scalariform perforations with ly diffuse-porous in wide growth rings (Fig. more bars (3-10) often present in the earliest 16). Vessels very abundant, usually more formed secondary xylem (Fig. 22). End walls than 200/mm2, sometimes up to more than of vessels oblique.