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Home , Achlys (plant), Berberis, Epimedium, Mahonia, Nandina, Podophyllum

PHYLOGENY AND WOOD ANATOMY OF

by

YU-FENGSHEN

Nqndina is a monotypic of great botanical interest. Its only , Nan­ dina domestica Tbunb. is an upright of central . It is commonly cultivated in China, known as Tien-chu (~~) or Heavenly , and is chiefly valued for its evergreen graceful foliage and its large panicle of bright red . Taxonomically it is of disputable position. It is generally included in the Berberi­ daceae. Aside from a few herbaceous genera, such as , jejJersonia, , , the only woody genera of the family are , , and Nandina. These genera are sometimes considered as constituting two subfamilies: Podophylloideae which comprise the above mentioned herbaceous genera and Berberi­ oideae, which comprise Berberis, Malwnia, Nandina as well as other herbaceous genera like and Le01ltice (Rendle 1925). The two subfamilies are considered by some authers as representing distinct families (Kumazawa 1930, 1938). Berberis and Mahonia are closely related. Actually they differ from each other only in the being always simple in the former and simple pinnate in the latter. Some authors express the opinion that the two should be treated as a single genus. Among all berberidaceous genera, the genus Nandina is .generally considered as deviate farthest phylogenetically. The aerial stem of this elongates monopodially until it terminates in as , producing two to four foliar and one to three scales each year. As the monopodial stem terminates in an inflorescence, the axillary bud elongates to form a new axis which in turn terminates in an inflorescence in the following year. The slender shrubby stem of the adult plant is, therefore, represented by a sympodial axis. (Kumazawa 1938). Kumazawa (1937) also investigates the ovular structures and types of dehiscence of the anther and finds that these characters are also unique among the berberidaceous genera. This genus, together with the other herbaceous genera such as EPimedium, Achlys, jejJersonia, etc. are included in the tribe Epimedieae by Prantl (1891) Tischler (1902), Himmelbaur (1913), and Engler-Diels (1936). Heintze (1927) establishes a Subfamily Nandinoideae for its sole accomodation and Nakai (1936; a family Nandinaceae. Ku· mazawa (1938) supports Nakai's view and considers it, together with Paeonia, as the most primitive members of the and Ranunculaceae. Various phases of morphological, anatomical, and biological studies on the Berberi­ daceae have been made by Citerne (1893), Tischler (1902), Himmelbaur (1913), Schmidt (1928), Champan (1936), Kumazawa (1936, 1937, 1938) and others. Record and Hess (1943), describe the wood anatomy of Berberis. . No investigation has been made on ;

86 the wood anatomy. of NaJldilla. As Xylotomy is now considered.as a helpful indicator of the phylogeny of woody , :the writer presents here the result of his investiga· tion on this problem. A description of the external characters of the three genera is as follows: . 1. Nandilla Thunb. Evergreen shrub: lvs. usually thrice pinnate; rachis articulate; Ifts. entire; tis. in large terminal panicles; in numerous whorls of 3, gradually passing into the white ; nectaries 3 or 6; anthers opening with longitudinal slit; a 2-seeded globular . 2. MallOnia Nutl. Evergreen unarmed , rarely small : terminal buds with numerous persistent p·ointed scales; leaves alternate. oddpinnate. rarely 3-foliolate; Ifts. usually spinose·dentate, the lat~ral sessile; stipules minute, subulate; tis. yellow in usually many·tid. or panicles springing from the axils of the bud·scales; .sepals 9; petals 6; ovary usually with few ovules; fr. dark blue and bloomy, rarely red or whitesh. About 50 species in North and Central Am. and in East and South Asia. 3. Berberis L. Evergreen or spiny shrubs rarely small trees: lvs. alter· nate, simple, those of the changed into usually 3-parted spines, the normal Ivs. appearing in fascicles on short axillary SpUTS; tis. yellow in elongated or umbel·like, rarely compound racemes, or solitary or fascicled; sepals 6, with 2 or 3 bracttets below; petals 6, often a smaller than the sepals and usually with 2 nectariferous glands at , base; 6, irritable, the anthers opening with 2 valves; ovary with 1 to many ovules; stigma sessile or on a short style; fro a 1- to several·seeded red to hlack berry. About 175 species, particularly well developed in E. and C. Asia and S. Am., few in N. Am., Eu. and N. Afr. For a study of the w90dy anatomy, radial, tangential, and transverse sections of the stem of Na",lilla domestica Thunb. are prepared from fresh material. Similar Sections of the stems of Berberis Kawakamii Hay. and Mahonia ~ommarifolia Takeda ar·o examined in order to corroborate Record and Hess's description (1943). They can· sid'~r the genus Berberis as including M~h'}nia and give the following description for the wood anatomy: "Pores small to minute, the large one zonate (ring·porous), the others arranged in an irregular pattern of wavy and zig.zag bands and patches. Vessels with simple perforations; members storied; spirals present; pitting fine, alternate. Rays nearly all large (Platnus type), giving ris~ to conspicuous silver grain on radial surface; not storied; nearly homogeneous; pits to vessel small, circular. Wood parenchyma sparse or absent. Wood fibers small, thick walled; pits very small. Ripple marks fairly distinct under lens". Their description fits very well the observation made by the present writer. Fur· thermore, the wood anatomy of Berberis and Mahonia does not reveal any essential differ~nces between the two' genera. The following is a description of the minute anatomy of the wood of Nalldina as studied by the present writer. Growth ring forming a semi·ring to diffuse porous condition. 87

Pore very smail (5 to 27 micro" in diameter), polygonal; numerous but not crowded laterally (48 to 56 per square millimeter); Pore wall thin, mostly 2-3 micron in thick­ ness; pores solitary or in radial multiples of 2 to 4. Vessel member storied, 184 to 328 micron in length; perforation plates simple, oblique. Intervessel pit-pairs bordered. very smal, 3.5 to 4 micron in diameter; alternate; distance between each pit l~terally 2 to 3. micron. Spiral thickening present in the inner wall of the vessel; tyloses thin-walled. Parenchyma terminal, metatracheal-diffuse, and paratracheal. Vvood rays, simple, aggregate, to oak type; not storied; homogeneous and hetero­ geneous; 1 to multi-seriate; 1 to 14 cells in thickness; 4 to many cells in height; crys­ tals absent; number" of rays per millimetre tangentially 2 to 5. Wood fibers with average length of 400--600 micron, middle diameter average 12 micron and with a wall thickness of 4--5 micron. Fiber long, tappering, sometimes with septate. Pits bordered, 2 to 3 micron in diameter; with lenticula to slitlike aper­ tures. The characters of the minute anatomy of the wood of M"Cllwnz"a and Berberz"s are very similar, but they differ from Nandina on the following three m~in accounts;- 1. Topography of wood. 2. Wood parenchyma. 3. Wood rays. The differences are as follows: (1) Nandina has diffused porosity, while Mahonia ~nd Berberis are all ring porous. (2) Nandina has more wood parenchyma; Berberis and'Mahonia has only little amount or even absent of parenchyma in the wood. ( 3) Nandina has very broad aggregate rays, mixed with parenchyma and fibers, while the rays of Mahonia and Berberz"s are simple and nearly homogeneouns. In "iew of these wide diff~rences, it may be concluded that in the wood anatomy Nandina is only remotely related to Berberz"s and MahlJnia. There differences are so great, that they seldom warrant to be included in the same family. Nandina differs greatly from Berberis and Mahonia in general external morphology. They also differ to a very large extent, as shown by the studies of various botanists mentioned above, in pollen morphology, ovular structure, and other minute anatomical features. The present author suports this view from a study of the wood anatomy and considers that Heintze (1927) and Nakai's (1936) proposales of establishing a subfamily Nandinoideae or a family Nandinaceae are worth considering.

LITERATURE CITED

(1) BAILEY, I. W. (1944): The development of vessels in angiosperms and its significance in morphological research. Arne!". Jour. Bot. 31: 421-428. (2) B--\RGHOORN, E. S., Jr. (1941): The ontogenetic development and phylogenetic specialization of rays in the Xylem of dicotyledons. I. The primitive ray structure. Amer. Jour. Bot. 27: 918-928. (3) '---, (1941a): The ontogenetic development and phylogenetic specialization of rays in 88

the Xylem of dicotyledons. n. Modification of the multiseriate and uniseriate rays. Amer. Jour. Bot. 28: 273-282. I (4) ---, (l941b): The ontogenetic development and phylogenetic speciation of rays in the Xylem of dicotyledons. III. The elimination of rays. Torrey Bot. Club. Bul. 68: 317-325. (5) CHALK, L. (1937): The phylogenetic value of certain anatomical tea,tures of dicotyledonous woods. Ann. Bot. 1: 409-428. ( 6 ) CHALK, L. and M. M. CHATTAWAY. (1934): Measur.ing the length of vessel members. Trop. Woods 1934 (40): 19-26. .. (7) CHAPMAN, M. (1936): Carpel anatomy of the Berberidaceae. Bot. Gaz. 23. (8) CITER~. P. E. (1893): Berberidees. et Erythrospennees. These Paris. (9) Committee on Nomenclature, International Association of Wood Anatomists. Glossary of tenns used in describing woods. Trap. Woods 1933 (36): 1-12.1933. (10) ENGLER, A. und L. DIELS (1936): Syllabus der Pflanzenfamilion. Autl. 11. Berlin. (11 ) Gilbert, S. G. (1940): Evolutionary significance of ~ing porosity in woody angiosperms. Bot. Gaz. 102: 105-120. ( 12) HANDLEY, W. R. C. (1936): Some observations on the problem of vessel length determination in woody dicotyledons. New Phytol. 35: 456-471. (13) HEINTZE, A. (1927): Cormofyternas fylogeni. lund. (14) HI.\ofl\1ELBAUR, W. (1931): Die Berberidaceen und ihre Stellung im System. Eine phyloge· netische Studie. Denkschr. dO. Wiss. math.-naturw. Klasse, 89. (IS) Ku~tAZAWA, M. (1936): Pollen grain morphology in Ranunculaceae, Lardizabalaceae and Berberidaceae. J~p. Jour: ~t. 8. (1'6) ---, (1937): Comparative studies on the vernation in the Ranunculaceae and Berberi· daceae. Jour. lap. Bot. 13. (17) ---, (1938 a): On the ovular structure in the Ranunculaceae and Berberidaceae. Jour. Jap. Bot. 14. (18) ---, (1938 b): Systematic and Phylogenetic Consideration of the Ranunculaceae and Berberidaceae. Bot. Mag. Tokyo LII: 613, page. 9-15. (19) LANGLE1.', O. F. (1928): Einige Beobachtungen Uber die Zytoiogie del' Berberidaceae Svensk. Bot. Tidskr. 22. (20) NA"KAI, ·T. (1936): Flora sylvatlca Koreana. 21. (21) PRANTL, K. (1891): Ranunculaceae, Berberidaceae. Englers natiirl. Pflanzenfam. Teil 3, Ab'.2. (22) RENDLE, A. B. (1925): The Classification of Flowering Plants, Vol. n. Dicotyledons. (23) RECORD, S.]. and HESS, R. W. (1943): Timbers of the New World. (24) SCHMIDT, E. (1928): Untersuchngen Uber Berberidaceen. Bcih. Bot. Centralbl. Abt. 2. 45. (25) TIPPO, O. (1946): The role of wood anatomy in pbylogeny. Amer. Midland Nat. 36: 362­ 372.

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O.5mm. II scale for the wood sections

Plate 1. Naftdiua domestica Thunb. A, of Nandina. B, wood of Nandina in radial section. C, wood of Nandina in tangential section. Multiseriate ray 'dissected by change of ray initials into fusiform initials. D, wood of Nandina in transverse se­ ction. with diffuced porosity. 90

Plate II. Mahollia /o11lmarijolia Takeda. A, shoot of Mahonia. B, wood of Mahonia in radial section. C, wood of Mahonia in tangential section. D, wood of Mahonia in transverse section. 9.1

Plate III. Buberis J(atvakamii Hay. A, shoot of Berberis. B, wood of Berberis in radial section. C, wood of Berberis in tangential section. D, wood of Berberis in tran· sverse section.