IAWA Bulletin n.s. , Vol. 13 (1),1992: 21-91 WOOD ANATOMY OF TREES AND SHRUBS FROM CIDNA. m. ROSACEAEI by Shu-Ym Zhang and Pieter Baas Rijksherbarium/Hortus Botanicus, P.O. Box 9514, 2300 RA Leiden , The Nether1ands Summary The wood anatomy of 162 species from lus, Armeniaca, Prunus S.str., Eriobotrya, China, belonging to 30 genera of the Rosa­ and Chaenomeles), as ornamentals (Rosa , ceae is described. The structural diversity Kerria, Spiraea, Sorbaria, Pyracantha, Photi­ is documented in a survey of characters, a nia, Exochorda, Docynia, and Chaenomeles), family description, generic descriptions and and as a source of valuable timber (Cotone­ tables. A key to the genera or groups of gen­ aster, Raphiolepis, and Pygeum). Fifty-four era is presented. A number of genera is de­ genera, about half of the total number ofgen­ scribed wood anatomically for the first time. era in the farnily, occur in China. In the pres­ Vestured pits noted in some Spiraea species ent study, thirty woody genera (see Table 1), are newly recorded for the farnily. The phe­ covering most of the woody ones, were sur­ nomenon of fibre dimorphism in Spiraea is veyed. A few small genera could not be in­ analysed in detail. cluded in the study because no wood sampies The systematic implications of the wood could be obtained. The classification of the anatomical diversity summarised in Tables family is subject to continued debate. For 22-24 are discussed at and below the sub­ practical reasons, we follow the systematic farnily level. The Spiraeoideae and Rosoideae arrangement and delimitation of subfamilies resemble each other very closely, but Exo­ and genera (see Table 1), and the nomencla­ chorda, and to a lesser extent Sorbaria, are ture as given in Flora Reipublicae Popularis aberrant within this group. Exochorda resem­ Sinicae (Yu 1974, 1985, 1986) although it is bles the Prunoideae in its wood anatomy. realised that the narrow generic and species The Maloideae constitute a very homogen­ concepts adopted are controversial. eous alliance, of which individual genera Wood anatomical data on Rosaceae in the cannot be separated, but two groups can be literature are mostly confined to certain gen­ recognised, only differing in degree of ray era of economic importance, and usually heterogeneity. The Prunoideae are wood ana­ scattered in limited papers and some books tomically the most heterogeneous and eight on wood anatomy of restricted regions. The groups in Chinese Prunoideae can be recog­ older literature was summarised by Solereder nised: Prinsepia and seven groups within the (1899, 1908) and Metcalfe and Chalk (1950). Prunus alliance (fable 24). More recent papers containing information Key words : Rosaceae, Spiraeoideae, Rosoi ­ on the wood anatomy of Rosaceae are: Baas deae, Maloideae, Prunoideae, comparative (1973), Baas et al. (1983, 1984), Baas & wood anatomy, wood identification. Schweingruber (1987), Barajas Morales (1980) , Brazier & Franklin (1961) , Burgess Introduction (1966), Van der Burgh (1978) , Bykova The Rosaceae are a farnily oftrees, shrubs (1966), Car1quist (1985, 1986, 1988a, b), and herbs ofcosmopolitan distribution. Many Carlquist & Hoekman (1985) , Cevallos-Fer­ genera in the family are of economic impor­ riz & Stockey (1990) , Cheng (1980, 1985), tance as fruit trees (Malus, Pyrus, Amygda- Cheng et al . (1979), Cheng & Liu (1991), 1) Dedicated to Prof. Dr. C. Kalkman, a specialist of the Rosaceae, on the occasion of his retirement as director of the Rijksherbarium/Hortus Botanicus. Downloaded from Brill.com10/09/2021 05:06:33PM via free access 22 IAWA Bulletin n.s., Vol. 13 (1),1992 Table 1. Enumeration of Chinese genera of the Rosaceae.* l".l l".l . ~ .~ 'ö 'ö 1l... .S'" 1l... .S'" e-c Subfamily I e-c Subfamily I "u "U Genera z .S Genera z .S Spiraeoideae (Rosoideaecontinued) Spiraea Shrub 50 10 Rubus Shrub-Tree 194 6 Sibiraea Shrub 3 Dryas Semi-shrub 1 Aruncus Herb 2 Geum Herb 3 Sorbaria Shrub 4 3 Acomastylis Herb 2 Physocarpus Shrub 1 Taihangia Herb 1 Neillia Shrub 10 Coluria Herb 3 Stephanandra Shrub 2 Waldsteinia Herb 1 Exochorda Shrub 3 Potentilla Herb 83 COmiJrum Herb 2 Maloideae Sibbaldia Herb 15 Dichotomanthes Shrub-Tree 1 1 Chamaerhodos Herb 5 Cotoneaster Shrub-Tree 58 3 Fragaria Herb 9 Pyracantha Shrub-Tree 7 1 Duchesnea Herb 2 Crataegus Shrub-Tree 17 9 Rosa Shrub 82 8 Osteomeies Shrub 3 Potaninia Shrub 1 Stranvaesia Tree-Shrub 4 1 Agrimonia Herb 4 Photinia Tree-Shrub 40 9 Spenceria Herb 1 Eriobotrya Tree-Shrub 13 4 Sanguisorba Herb 7 Raphiolepis Shrub-Tree 7 4 Alchemilla Herb 3 Sorbus Tree-Shrub 55 25 Cydonia Shrub-Tree 1 1 Prunoideae Docynia Tree 2 1 Prinsepia Shrub 4 2 Chaenomeles Shrub-Tree 5 3 Amygdalus Tree-Shrub 12 5 Pyrus Tree-Shrub 14 8 Armeniaca Tree 7 6 Malus Tree-Shrub 22 14 Prunus s.str. Tree-Shrub 7 4 Amelanchier Shrub-Tree 2 1 Cerasus Tree-Shrub 45 15 Rosoideae Padus Tree-Shrub 14 9 Kerria Shrub 1 Laurocerasus Tree-Shrub 13 6 Rhodotypos Shrub 1 Pygeum Tree-Shrub 6 1 Filipendula Herb 8 Maddenia Tree-Shrub 5 * Based on the Flora Reipublicae Popularis Sinicae (Yu 1974, 1985, 1986). Chudnoff (1956), Cristiani (1962), Cutler er Huang (1964), Huber & Rouschal (1954), al. (1987), Dechamps (1985), Desch (1954), Jacquiot er a/. (1973), Jagiella & Kurschner Detienne & Jacquet (1983), Detienne er al. (1987), Janssonius (1952), Kribs (1968), (1982), Duperon (1976), Fabbri Tarchi Lebacq (1957), Luo (1989), Meylan & But­ (1960, 1963), Fahn er al. (1986), De Freitas terfield (1975), Niloufari (1961), Normand (1958), Friedman (1978), Furuno (1979, (1950), Novruzova (1962, 1964), Novru­ 1985), Gabrieljan (1954, 1971), Grambast­ zova & Gadzhieva (1974), Ogata (1975­ Fessard (1966), Greguss (1959), Grosser 1983), Oprea (1972), Page (1964), Panshin & (1977), Guleria er al. (1983), Hayashi er a/. De Zeeuw (1980), Parsa Pajouh & Schwein­ (1973), Ho (1985), Hofmann (1944, 1955), gruber (1985) , Poller (1967), Prive-Gill Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 23 (1981), Rao & Purkayastha (1972), Schwein­ Zhang 1986; Baas et al. 1988; Deng & Baas grober (1974, 1978, 1990), Seimeier (1984), 1990). The definition of the anatomical fea­ Snezhkova (1977,1979), Sudo (1959,1963), tures used in the Rosaceae and measurements Süss & Müller-Stoll (1983), Suzuki (1984), of quantitative features largely follow these Suzuki et al. (1991), Takahashi & Suzuki papers and the IAWA List (Wheeler et al. (1988), Tang (1976), Tumanin (1949,1954), 1989). Tumanjan (1950) , Venet (1974), Wang The ratio of vessel element length/vessel (1965, 1966), Wheeler & Matten (1977), diameter (LID ratio) is used here as an index Wheeler et al. (1978), Wu et al. (1989), Yang of vessel element shape, rnainly because ves­ & Huang-Yang (1987), Yaskevich (1956) and sel element diameter and vessel element Yatsenko-Khmelevsky (1954). Most publica­ length in many genera vary considerably. tions listed above deal with species of the Thus the ratio gives additional information. subfamilies Maloideae and Prunoideae. Very few publications are concemed with repre­ sentatives of the Spiraeoideae and Rosoideae. Survey ofwood anatomical features in the Rosaceae from China In this study we will focus on a descrip­ tive survey and separation of genera and sub­ Introduction families in China as based on their wood anatomy. A worldwide survey exploring the The following survey describes the range systematic and phylogenetic wood anatomy of variation in various wood anatomical fea­ of the Rosaceae and a study of ecological tures in the Chinese Rosaceae. It also focuses on the diagnostic and systematic value of var­ trends are in progress. ious wood anatomical features, especially at the level of subfamilies or groups of presum­ Materials and Methods ably closely related genera. In addition, a More than 300 wood samples were obtain­ wood anatomical family description of Chi­ ed, mainly from various Chinese institutional nese Rosaceae is given. wood collections, complemented with field Growth rings (Figs. 1-12) collections of mainly shrubby species by sev­ Growth rings in the Chinese Rosaceae eral taxonomists (see Acknowledgements, vary from distinct in most genera studied to page 86). Many of the samples studied, un­ faint or even absent. The following growth fortunately, are not vouchered. However, as ring markers occur in various combinations: much as possible more than one samples (up 1) flattened latewood fibres (almost always ro nine) per species were studied to enhance present), 2) differences in vessel diameter the chance of spotting erroneous identifica­ and/or frequency between latewood and tion and to study infraspecific variation . For subsequent earlywood (semi-ring-porous and each sample, the locality is listed first, fol­ ring-porous wood), 3) marginal parenchyma lowed by the wood collection number pre­ bands, 4) wavy growth ring outline and/or ceded by a code from Stern's Index Xylari­ locally inflated rays. In the Maloideae and orum (1988), indicating the source of the most Prunoideae growth rings are only mark­ sample. For shrub or branch sarnples (e.g . ed by flattened latewood fibres, with the ex­ Spiraea and Rosa), sample diameter is given. ception of some ring-porous genera (e.g . Altitude is indicated ifinformation was avail­ Amygdalus & Armeniaca), and Pygeum & able. Laurocerasus Group B which have marginal All sarnples were sectioned and macerated parenchyma bands. More or less wavy boun­ according to the standard techniques described daries, sometimes associated with locally in­ by Baas and Zhang (1986) for light micro­ flated broad rays are typical of most genera in scopic study, followed by SEM observations the Spiraeoideae and Rosoideae. on certain features of all the samples, The sequence adopted for the generic de­ Vessels scriptions in the Rosaceae follow a standard Porosity (Figs.7-1O) - Ring-porous, forrnat presented in three papers (Baas & serni-ring-porous and diffuse-porous woods Downloaded from Brill.com10/09/2021 05:06:33PM via free access 24 IAWA Bulletin n.s., Vol.