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.

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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

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(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. 13 (1),1992 all occur in the family, even within a single Arrangement and grouping (Figs. 2, 8, subfamily, the Prunoideae for instance. Most 10-12) - Vessels mostly show a more or genera in the Chinese Rosaceae have diffuse­ less random distribution in the family. How­ porous woods (rarely ranging to semi-ring­ ever, vessels in an oblique to radial pattern porous woods), especially the Maloideae. were noted in some species of the Prunoideae Typical ring-porous woods are limited to a and to a lesser extent in Spiraea p.p. few genera like Rosa . In certain genera, po­ The percentage of solitary vessels is of rosity is variable. Thus two types and their some diagnostic value in the Rosaceae from intermediates can be recognised.such as dif­ China, despite large variation. With few ex­ fuse-porous to semi-ring-porous in Cerasus, ceptions, the percentage of solitary vessels Cotoneaster, Padus, Prinsepia, Spiraea and is less than 55(-60) % in all genera of the Stranvaesia, or ring-porous to semi-ring­ Prunoideae (except in Prinsepia), while it is porous in Exochorda. Within Prunus s.l. over (50-)55% up to 99% in the remaining (cf. Bentham & Hooker 1865) divided into three subfamilies (except in Rubus). The six genera by Yu (1986), Amygdalus and Ar­ non-solitary vessels are mainly in tangential meniaca are ring-porous, while the remaining and oblique (rarely radial) multiples of 2-4 genera are usually diffuse-porous, rarely (-5) in the Spiraeoideae and Rosoideae , in ranging to semi-ring-porous in some species of Cerasus, Padus and Prunus s.str. (text continued on page 38)

Legends to Figures 1-61:

Fig. 1. TS, x 73. Growth ring boundaries indistinct and vessels of two sizes in Rubus bif/orus. - Fig. 2. TS, x 46. Growth ring boundaries absent or indisrlnct in Pygeum topengii and with a high degree of vessel grouping. - Fig. 3. TS, x 46. Growth ring boundaries faint, marked by one row of discontinuous marginal parenchyma in Laurocerasus fordiana . - Fig. 4. TS, x 46. Growth ring boundaries more or less distinct, marked by 2-3-seriate marginal parenchyma bands in Laurocerasus hypotricha.

Fig. 5. TS, x 46. Growth ring boundaries marked by marginal parenchyma bands over 5 cells wide, mixed with traumatic gum ducts in Pygeum topengii . - Fig. 6. TS, x 46. Growth rings boundaries wavy, and marked by differences in vessel diameter between latewood and subse­ quent earlywood in Rosa macrophy//a. - Fig. 7. TS, x 46. Wood diffuse-porous, growth ring boundaries distinct, marked by 1-3 rows of radially flanened latewood fibres in Malus baccata. - Fig. 8. TS, x 46. Growth ring boundaries marked by locally inflated broad rays, vessels mostly solitary and the widest vessels occurring in intermediate position of growth rings in Sor­ baria arborea var. subtomentosa.

Fig. 9. TS, x 46. Wood ring-porous with pith flecks in Armeniaca vulgaris. - Fig. 10. TS, x 46. Wood semi-ring-porous and vessels in a weakly expressed radial pattern and mostly solitary in Spiraea thunbergii. - Fig. 11. TS, x 46. Vessels in a radial pattern and with a high degree of grouping in Laurocerasus zippe/iana. - Fig. 12. TS, x 46. Vessels in a weakly pro­ nounced oblique pattern and with a fairly high degree of grouping in Laurocerasus spinulosa.

Fig. 13. RLS, SEM, x 1080. Scalarifonn perforation plate with 15 bars in Malus doumeri.­ Fig. 14. RLS, SEM, x 3600. Irregular perforation plate in Sorbus commixta. - Fig. 15. RLS, SEM, x 648. Three irregular perforation plates in Photinia beauverdiana.- Fig. 16. RLS, SEM, x 2160. Irregular perforation plate in Sorbus pteridophy//a . - Fig. 17. RLS, SEM, x 2160. Irregular perforation plate in Photinia vi//osa. - Fig. 18. RLS, SEM, x 2340. Irregular perfo­ ration plate in Sorbus pteridophy//a .

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Fig. 19. RLS, x 443. Intervesse1 pits alternate, polygonal, apermres occasionally coalescent in Rosa macrophylla. - Fig. 20. TLS, x 273. Intervessel pits alternate, mostly polygonal, and helical thickenings closely spaced in Laurocerasuszippeliana. - Fig. 21. RLS, x 273. Vessel­ ray pits clearly smaller than intervessel pits in AmygdaJus triloba.- Fig. 22. RLS, x 443. Ves­ sel-ray pits in Chaenomelesthibetica. - Fig. 23. RLS, SEM, x 2160. Intervessel pits alternate, round with oval to slit-like apertures in Sorbus alnifolia.- Fig. 24. RLS, SEM, x 2340. Inner vessel wall showing coalescent pit apertures in Amygdalusdavidiana .

Fig. 25. RLS, SEM, x 117oo. Vestured pits in Spiraea thunbergii. - Fig. 26. RLS, SEM, x 8460. Weakly vestured pits in Spiraeajaponicavsx.fortunei.- Fig. 27. RLS, x 443; 'fused' in­ tervessel pits (arrow) in vessel wall of Raphiolepis indica.- Fig. 28. RLS, x 273. Vasicentric tracheids with distinctly bordered pits and irregular in shape in Armeniaca vuIgaris. - Fig. 29. RLS, SEM, x 350. Irregular thickenings associated with pit apertures in Sorbus hemsleyi. ­ Fig. 30. RLS, SEM, x 1800. Well-developed and closely spaced helical thickenings in Erio­ botryajaponica.- Fig. 31. RLS, SEM, x 900. Widely spaced thickenings in Padusbuergeriana.

Fig. 32. RLS, SEM, x 1800. Well-developed, coarse helical thickenings in Laurocerasushypo­ tricha.- Fig. 33. RLS, SEM, x 1350. Poorly developed, fine vessel wall thickenings in Pygeum topengii. - Fig. 34. RLS, SEM, x 1350. Irregular thickenings in Cerasusdielsiana.- Fig. 35. RLS, SEM, x 1980. Poorly developed irregular thickenings in Rosa cymosa.- Fig. 36. RLS, SEM, x 288. Tyloses in vessels in Padus obtusata. - Fig. 37. RLS, SEM, x 540. Gummy con­ tents in vessel in Armeniacaholosericea.

Fig. 38. RLS, x 443. Coarse helical thickenings in fibre-tracheids ofCrataegusoresbia. - Fig. 39. TLS, x 273. Fibre-tracheids with distinctly bordered pits in Sorbusprattii. - Fig. 40. RLS, SEM, x 1800. Libriform fibres with simple pits in Spiraea blumei. - Fig. 41. RLS, SEM, x 2160. Fibre-tracheids with distinctly bordered pits in Spiraea blwnei. - Fig. 42. RLS, SEM, x 1800. Libriform fibres (right) with simple to minutely bordered pits and fibre-tracheids (left) with distinctly bordered pits in Spiraea thunbergii. - Fig. 43. TS, x 110. Parenchyma mainly diffuse-in-aggregates and scanty paratraeheal, and pith fleck present in Photinia prunifolia.

Fig. 44. x 116. Fibre pits almost absent in tangential walls and intervessel pits alternate, aper­ tures usually coalescent in Pygeum topengii. - Fig. 45. TLS, x 46. Juvenilistic rays, of two distinct sizes, in Spiraea cantoniensis. - Fig. 46. TLS, x 46. Rays of two sizes, but clearly lower in Cerasus conradinae. - Fig. 47. x 46; 1-4-seriate rays in LaurocerasusspinuIosa.

Fig. 48. TLS, x 116; 1-2(-3)-seriate rays in Sorbus discolor.- Fig. 49. TLS, x 116. Uniseri­ ate rays in Crataegusoresbia. - Fig. 50. RLS, x 288. Rays composed ofprocumbent cells only in Crataegusshensiensis. - Fig. 51. x 185. Rays composed ofprocumbent body cells and 1-3 rows of square to upright cells in Dichotomanthes tristaniaecarpa.

Fig. 52. RLS, x 288. Perforated ray cell and rays composed of square to procumbent cells in SpiraeachamaedryJolia. - Fig. 53. TLS, x 116. Weakly differentiated sheath cells in juvenilis­ tie rays in Rubus chingii.- Fig. 54. TLS, x 288. Inflated secretory (?) cells (possibly oil or muci­ lage) associated with ray parenchyma in Cerasus campanulata.- Fig. 55. RLS, x 467. Pris­ matic crystals in chambered and enlarged parenchyma cells in Dichotomanthes tristaniaecarpa.

Fig. 56. RLS, x 110. Druses in ray cells in Cerasus setuIosa. - Fig. 57. RLS, x 273. Prismaric crystals in axial parenchyma cells, non-enlarged, usually one crystal (occasionally more than one) per cell in Spiraea chamaedryfolia. - Fig. 58. RLS, SEM, x 3600. Druse in ray eell in Cerasus setuIosa. - Fig. 59. RLS, SEM, x 2160. Prismatie crystals in ehambered and enlarged parenehyma eells in Malus specrabilis. - Fig. 60. RLS, x 110. Prismarie crystals in ehambered and enlarged axial parenchyrna cells, one crystal per chamber in Photiniaglabra. - Fig. 61. TS, x 43. Traumatic gum ducts in tangential arrangement in Armeniacavulgaris.

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Percentage of sampies (%) for the Prunoideae to have wider vessels than 60 ,------, the other three subfamilies. The Maloideae, Spiraeoideae and Rosoideae share almost the same range in tangential vessel diameter. For radial vessel diameter, more or less the same trends were noted . The most frequent range is from 30-80 11m. Solitary vessels are mainly round to oval in outline in the Spiraeoideae and Rosoideae. Apart from round to oval vessels, more or less angular vessels can be found in the Mal­ oideae and Prunoideae, except in Pygeum 10 20 30 40 50 60 70 80 90 100 110 120 Tangential vessel diameter (um) Percentage of sampies (%) Fig. 62. Tangential vessel diameter ranges in 50,------._------, the subfamilies.• = Spiraeoideae & Rosoi­ 45 deae; "* =Maloideae; ... =Prunoideae. 40 ;\\ I 35 30 25 oblique (rarely radial and tangential) pairs or short multiples in the Maloideae, or mainly in 20 oblique and/or radial multiples of 2-4(-12) 15 in the Prunoideae . Oblique to radial multiples 10 of over 4 vessels are common in some spe­ 5 eies of Padus and Laurocerasus, but occur O J...... ,-~~-~~I -_r_~-_r_~-.,...... J 100 200 300 400 ..500 600 700 800 900 1000 sporadically in the remaining genera of the Prunoideae. Vessel clusters of 3 to 4 occur Vessel element length (um) sporadically in a few species of Laurocerasus. Fig. 63. Vessel element length ranges in the subfamilies.• = Spiraeoideae & Rosoideae; Frequency anti element size (Figs. 62-64) "* =Maloideae; ... =Prunoideae. - Vessel frequency ranges from 8 vesselsl sq.mm in Pygeum topengii to 520 vessels/ Percentage of sampies (%) 35 ,------, sq.mm in Crataegusmaximowiczii.But most genera (or speeies) have an average vessel 30 frequency of 100-200 vessels/sq.mm. Ves­ seI frequency varies greatly within some gen­ 25 era (Spiraea and Amygda/us), while in some 20 other genera (e.g. Rubus and Photinia) it shows a comparatively narrow range . There 15 are no appreeiable differences in vessel fre­ quency range between the subfamilies. 10

Average tangential vessel diameter ranges 5 from 16 um in Raphiolepis indicato 123 um in Pygeum topengii. It is, however, less than 60 um in most genera studied. Figure 62 o 2 4 6 8 10 12 14 16 18 2022 24 26 28 30 shows the frequency distribu tion of average LID ratio tangential vessel diameter in the subfamilies. Fig. 64. LID (ratio of vessel element length The most frequent range for tangential diam­ and diameter) ranges in the subfamilies.• = eter in the Chinese Rosaceae is from 20-70 Spiraeoideae & Rosoideae; "* = Maloideae; um. Despite large overlap, there is a tendency ... = Prunoideae.

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 39 and Group B of Laurocerasus. The ratio of ous irregular multiple perforations occur in radial diameter/tangential diameter ranges on about half of the genera studied: Amygdalus, average from 1.15-1.55. The Spiraeoideae Cerasus, Crataegus, Eriobotrya, Kerria, Ma­ and Rosoideae tend to have a slightly lower lus, Padus, Photinia, Pyracantha, Pyrus, ratio (1.15-1.30) than the Maloideae (1.20­ Raphiolepis, Rubus, Sorbus, Spiraea, Steph­ 1.55). anandra. This list covers all four subfarnilies. Vessel element length also shows a wide Usually scalariform and/or irregular perfora­ range from 230 um in Armeniacavulgaris to tions occur in very few species of these gen­ 930 um in Eriobotrya cavaleriei. As shown in era, and were sometimes only found in one Figure 63, vessel element length distributions species (e.g. Rubus and Amygdalus) or one in the Prunoideae , Spiraeoideae and Rosoi­ sample only (e.s; Stephanandra and Kerria). deae are almost the same,ranging from (200-) Furthermore, the frequency of scalariform 300-600(-700) um, But the Maloideae tend and/or irregular perforations is very low to have Ionger vessel elements with a range (usually 1-5%). Sometimes only one mul­ of 300-900 um. Thus the Prunoideae gener­ tiple perforation can be found in one slide. ally have wider but shorter vessel elements Therefore it is uncertain whether the perfora­ than the Maloideae. In other words, the ratio tions are exclusively simple in the remaining of vessel element length/vessel diameter genera. Scalariform and/or irregular perfora­ (LID ratio) in the Prunoideae is lower, with tions probably could have been found in a full range of 4-14(-18), as shown in Fig­ more genera if more species and samples had ure 64. Among them, Pygeum has the lowest been available. In the Maloideae, multiple LID ratio (4-5), followed by Group B of perforations are slightly less infrequent than Laurocerasus (5-7). In all other genera of in the other subfarnilies, and of relatively the Prunoideae , however, the LID ratio common occurrence in Sorbus, Eriobotrya, mostly ranges from 5 to 14. The Maloideae Raphiolepis, Malus and Pyrus. The number stand out on account of a high ratio, mostly of bars mostly ranges from 2 to 5, rarely up ranging from 10-24 (Fig. 64). The highest to 15. The perforations include scalariform, ratio occurs in Raphiolepis, Eriobotrya and reticulate, foraminate and various other irreg­ Sorbus. The Spiraeoideae and Rosoideae ular types (see Figs. 13-18). have a range of (6-)8-18. But Sorbaria is exceptional with a low ratio (6-8), even Wall pitting (Figs. 19-27) - Intervessel lower than most species in the Prunoideae. pits are typically alternate, but in addition Part of the variation recorded here for ele­ scalariform to opposite pits were also noted ment dimensions is probably related to large in Kerria. Intervessel pit shape tends to differ differences in plant habit (ranging from small from subfamily to subfarnily: round to oval in shrubs to big trees) and to non-standard sam­ the Maloideae; mainly polygonal in the pling (branch wood versus stern wood; vary­ Prunoideae, especially pronounced in Cera­ ing height in the stern; stern age and diameter, sus, Padus, Laurocerasus and Pygeum; inter­ etc.). The small stature of most Spiraeoideae vessel pit shape is variable in the Spiraeoideae and Rosoideae thus may partly account for and the Rosoideae. Round, oval and polygo­ the low values for vessel size and fibre length, nal pits are all represented. Pit apertures are and high values for vessel frequency. mainly oval to slit-like. Coalescent apertures Vessel wall thickness ranges from 1-2(-3) are occasionally present in some species. um in most genera. Pygeum and Group B of Intervessel pits are usually nonvestured, but Laurocerasus species are exceptional with SEM studies revealed that vestured pits occur walls of 2-4lUIl thick. in five Outof ten species studied in Spiraea. The vestured pits are mainly restricted to Perforation plates (Figs. 13-18) - Rosa­ the regions near the perforation rims. Bailey ceae typically have simple perforations, main­ (1933) reported that vestured pits are absent ly in oblique, more rarely in (nearly) hori­ in Rosaceae, and that the sieve-like structures zontal end walls depending on vessel shape in Cerasusserotinusand Prunus brigantiaca (LID ratio). Sporadic scalariform and vari- referred to by Jönsson (1892) are probably

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artifacts. The recognition of vestured pits in ments in a few genera, or confined to narrow Spiraea therefore is a new record for Rosa­ vessel elements or even to the tails of narrow ceae. Vestured pits are generaily assumed to elements only. Most genera of the Prunoideae be of high diagnostic and systematic value are characterised by weil developed helical (Bailey 1933). However, the vestured pits in thickenings. The thickeningsare usually wide­ Spiraea are exceptional for their sporadic dis­ ly spaced, coarse and present throughout the tribution, and thus of limited diagnostic and vessel elements in Amygdalus, Armeniaca, systematic value. Prunus, Cerasus, Padusand Group A of Lau­ Intervessel pit size varies appreciably in rocerasus. However, vessel wall thickenings the farnily, ranging from 2-12(-20) um with in Pygeumand Group B of Laurocerasus are the ranges for individual subfarnilies largely closely spaced, sometimes less developed or overlapping. Kerria is unique for its large confined to narrow vessel elements or to the range of intervessel pit size (3-20 um) relat­ tails of narrow vessel elements only. ed to the presence of alternate to scalariform Contents and tyloses (Figs. 36, 37) ­ (to opposite) pits. Pygeum and Group B of Gummy contents are common in the Prunoi­ Laurocerasus have intervessel pits of2-4(-5) deae, occasionally present in the Maloideae um in diameter, significantly smaller than in and almost always absent in the Spiraeoideae the other genera of the Prunoideae. and Rosoideae due to the absence of heart­ Vessel-ray and vessel-parenchyma pits wood formation in thin sterns of shrubby spe­ are also variable in size. In most genera of cies. Tyloses are almost always absent in the the Spiraeoideae, Rosoideae and Maloideae, farnily, and were only noted in few species. the pits are more or less similar to the inter­ vessel pits but half-bordered and/or a little smaller. However, vessel-ray and vessel­ Vasicentric tracheids (Fig.28) parenchyma pits are usually much smaller Vasicentric tracheids are restricted to Rosa thanthe intervessel pits and have reduced bor­ and a few species in the Prunoideae. Vasicen­ ders in most Prunoideae, except in Pygeum tric tracheids are mainly present in earlywood and Group B of Laurocerasus where vessel­ and irregular in shape. They have distinctly ray and vessel-parenchyma pits are sirnilar to bordered pits in both radial and tangential the intervessel pits. walls.

Helical and other wall thickenings (Figs. Ground tissue fibres (Figs. 38-42, 44, 65) 29-35) - Vessel wall thickenings are usual­ The Maloideae are characterised by ground ly absent in Spiraeoideae except in Exochor­ tissue fibres with distinctly bordered pits of da where closely spaced thickenings occur, mostly 4-7 (3-9) um in diameter, common mainly confined to narrow vessel elements. in radial and tangential walls. In other words, The same applies to the Rosoideae where the ground tissue in the subfarnily is compos­ vessel wall thickenings are usually absent ed exclusively of fibre-tracheids as defined (e.g., Kerria and Rubus), but irregular and by Baas (1985, 1986). In the Spiraeoideae discontinuous thickenings are of constant (except Spiraea) and the Rosoideae, the occurrence in Rosa (confined to the body of ground tissue is also composed of fibre­ narrow vessel elements). In the Maloideae, tracheids with distinctly bordered pits, but the the feature varies considerably: e.g. helical pits are mainly concentrated in the radial thickenings constant in Amelanchier, Coton­ walls. The diameter of the pits in these two easter, Cydonia,Dichotomanthus, Docynia, subfarnilies mostly ranges from 3-5(-7) the deciduous group of Photinia,Pyracantha, um. Spiraeais unique for having two diverse Raphiolepis, Sorbus and Stranvaesia , or re­ types of ground tissue fibres: libriform fibres stricted to some species in Chaenomeles, Cra­ with simple to minutely bordered pits of 1-2 taegus, Eriobotrya, Malus, the evergreen um and fibre-tracheids with distinctly border­ group of Photinia, and Pyrus . The wall thick­ ed pits of 3(-4) um in diameter. In the Prun­ enings in the Maloideae are usually closely oideae, the ground tissue fibres vary consid­ spaced, present throughout the vessel ele- erably, ranging from libriform fibres with

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Percentage of sampIes (%) As shown in Figure 65, the ratio of fibre 45 -r------, length to vessel element length (FIV ratio) in 40 the Prunoideae , mostly 1.8-3.4, tends to be higher than in the other three subfamilies 35 which share about the same range of 1.4-2.6. 30 Helical thickenings in ground tissue fibres 25 are usually absent in the Spiraeoideae, Rosoi­

20 deae and Prunoideae. In the Maloideae , heli­ cal fibre wall thickenings vary from genus to 15 genus. They are of constant occurrence in all 10 Chinese species of some genera (Cotoneaster, Crateagus, Raphiolepis and Chaenome/es ); restricted to some species (Photinia, Sorbus 1.2 1.4 1.6 1.8 2 2.2 2.4 2.6 2.8 3 3.2 3.4 3.6 3.8 4 and Pyrus); or absent (Eriobotrya and Malus). F/V ratio When present, the thickenings are usually Fig. 65. FIV (ratio of fibre to vessel element weil developed in some to almost all fibre­ length) ranges in the subfamilies . • = Spir­ tracheids. Ground tissue fibres are almost always aeoideae & Rosoideae ; * = Maloideae; ... = Prunoideae. nonseptate in the Rosaceae from China. Sporadic septate fibres were noted in a few species of Spiraea.

Axial parenchyma (Figs. 1-12,43) simple to minutely bordered pits of 1-2 um, Both types and amount ofaxial parenchy­ to fibre-tracheids with distinctly bordered pits ma are variable in the family. The following of 3-5 IJ.m. Most species in the subfamily types of parenchyrna distribution occur in the have fibres with distinctly bordered pits of Chinese Rosaceae : 1) apotracheally diffuse; 3-5 IJ.m, mainly confined to the radial walls. 2) scanty paratracheal; 3) diffuse-in-aggre­ Generally fibre pits in this subfamily are less gates; 4) marginal bands and/or narrow ir­ distinctly bordered than in the Maloideae. regular zonate bands. Most genera usually Typicallibriform fibres in the Prunoideae are have at least two of the above types. Apotra­ restricted to a few species (e.g., Amygdalus cheally diffuse and scanty paratracheal paren­ davidiana, A. persica, Cerasus maximowiczii chyma are typical for most Spiraeoideae and and C. yedoensis). Rosoideae where the amount ofaxial paren­ The walls of ground tissue fibres are me­ chyma is usually scanty. However, Exochor­ dium thick to very thick in most genera, but da and Sorbaria are characterised by more thin-walled fibres are also present in a few abundant parenchyma and here the diffuse­ genera like Spiraea and Pygeum. This means in-aggregates type is also present. In the that most genera probably have a medium or Maloideae, axial parenchyma is relatively relatively high wood density, which is con­ abundant, and thus diffuse-in-aggregates pa­ firmed by the limited data available (Cheng renchyma is common in addition to apotra­ 1985; Huang 1964). cheally diffuse and scanty paratracheal paren­ Like other quantitative features, length of chyma. Axial parenchyma in the Prunoideae ground tissue fibres varies greatly, from is variable in both amount and arrangement: 400-1600 IJ.m. There is a tendency for the usually scanty diffuse and paratracheal in Spiraeoideae and Rosoideae to have shorter most genera, but Pygeum and group B of fibres, ranging from 400-1100 IJ.m . Fibre Laurocerasus have more abundant parenchy­ length in the Prunoideae and Maloideae is ma, including marginal parenchyma bands. more or less the same, ranging from 600­ Axial parenchyma strands are usually 3-5 1500 IJ.m. Spiraea is characterised by the cells long, with a total range of 2 to 11 cells. shortest ground tissue fibres with average There are no differences between the sub­ values ranging from 400-600 IJ.m. families.

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Ray tissue (Figs. 44-53) Crystals (Figs. 55-60) In the Rosaceae from China, rays vary Crystals are not of constant occurrence in tremendously in size and composition, par­ most genera. Occurrence, type, size, relative ticularly at the level of the subfamilies. The abundance and the type of cells containing the Spiraeoideae and Rosoideae are mostly char­ crystals all may vary. Crystals are present in acterised by typical juvenilistic rays. The rays some species of the Prunoideae and Spiraeoi­ are composed of square or square to procum­ deae & Rosoideae, but they are more fre­ bent body cells and upright marginal cells, quent in the Maloideae. Two types of crys­ usually also with weakly differentiated sheath tals, prismatic and druses occur. Prlsmatic cells. The rays in these two subfamilies are crystals are the more common type, present of two distinct sizes: narrow and low rays, in all four subfamilies. Druses are restricted usually 1(-2)-seriate, and wide and tall rays, to the subfamily Prunoideae. The crystals in more than (3-)4-seriate up to Iö-seriate. The Spireaoideae and Rosoideae are sparse to wide and tall rays are higher than 1.0 mm, up common, medium-sized (occasionally mi­ to 25.0 mm, and the widest rays are usually nute), and almost always present in non­ over ö-seriate up to 16-seriate. Exochorda chambered and non-enlarged ray cells (except and Sorbaria in the Spiraeoideae are excep­ in Spiraea where rare crystals are also found tional: their rays are lower than 1.0 mm and in chambered but non-enlarged axial paren­ body ray cells are procumbent. Rays of two chyma cells of one species). In the Maloideae, sizes can be recognised in most genera of however, prismatic crystals are sparse or the Prunoideae , but they are not as distinct common to abundant, usually large, and re­ as in the Spiraeoideae and Rosoideae . The stricted to chambered axial parenchyma cells multiseriate rays are lower than 1.0 mm, but (or sometimes to idioblasts) . Each enlarged higher than 0.3 mm, and narrower with a and sclerified chamber (or idioblast) contains range of 4-11-seriate for the widest rays. one crystal . In the Prunoideae, the crystal The multiseriate rays in the Prunoideae are complement is variable: druses and/or pris­ composed of procumbent body cells and matic crystals are present in ray cells and/or 1-5(-7) rows of square to upright marginal axial parenchyma cells, or crystals are com­ cells in most genera (except in Padus). In pletely absent. The ray cells containing crys­ the Maloideae, rays are of intergrading sizes tals in the Prunoideae are usually non-cham­ only, mostly 1-2(-3)-seriate, and the aver­ bered and non-enlarged, but the reverse holds age height of the multiseriate rays is not more true for the axial parenchyma cells, especially than 0.4 mm. The multiseriate rays are com­ in Cerasus. Crystals have not been observed posed of procumbent cells only, or some­ in Prunus s.str. and Pygeum. However, times have one row of square marginal cells crystals were found in almost all species of in some genera (Amelanchier, Crataegus, Cerasus. Cydonia, Docynia, Malus , Pyrus and Sor­ bus), or are composed of procumbent body Pith flecks (Figs. 8, 9, 43) cells and 1-6 rows of square to upright mar­ Pith flecks are of sporadie occurrence in ginal cells in the remaining genera (Chaeno­ all four subfamilies and are not of diagnostic meles , Cotoneaster, Dichotomanthes, Erio­ value. They are more common in the Maloi­ botrya, Photinia, Pyracantha,Raphiolepis and deae and Prunoideae than in the other two sub­ Stranvaesia). families. Pith flecks are diffuse or arranged in Ray frequency varies greatly with a range tangential bands. of 5-17/mm, with overlapping ranges for all subfamilies. Traumatic gum ducts (Figs. 5, 61) Perforated ray cells were observed only in Traumatic gum ducts are restricted to the Spiraea chamaedryJolia. Large secretory (?) subfamily Prunoideae . They were observed cells (with unidentified contents, possibly in a few Chinese species of all genera except oil or mucilage)associated with ray paren­ in Prunus s.str. However, the ducts are vari­ chyma were noted in Cerasus campanulata able within a genus or even a single species. (Fig. 54). They may be present in one sample but be

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 43 absent in another sampIe of the same species tured except in some Spiraea species, usually (in Amygdalus persica for instance). In size alternate, round, oval and polygonal, 2-12 and position in a growth ring, the traumatic (-20) J.LIIl in diameter, with slit-like apertures, gum ducts are more or less similar to the pith apertures occasionally coalescent. Vessel-ray flecks . Usually traumatic gum ducts are ar­ and vessel-puenchyma pits mostly similar to ranged in short to long tangentiallines, some­ intervessel pits but usually half-bordered and times they are diffuse. The ducts occur slightly smaller, or elearly smaller than inter­ mainly in earlywood or in the intermediate vessei pits in a few genera. Helical or irregu­ wood of growth rings . lar thickenings present in many genera, elose­ ly to widely spaced, fine to coarse, through­ Summary and fiunily description ofthe out all vessel elements or restricted to narrow Chinese Rosaceae vessel elements (or to the tails only) . Tyloses Based on the above survey of wood ana­ almost always absent. Gummy contents pres­ tomical features in the Chinese Rosaceae, the ent or absent. following features, to differing extent, appear Vasicentric tracheids, usually ofirregular to be of some taxonomic value mainly ar sub­ shape, sparsely occur in some species of a family level or to characterise groups of pre­ few genera only, mainly in earlywood. sumably elosely related genera: I) vessels Ground tissue fibres 400-1600 um long, (degree of vessel grouping and vessel wall F/V ratio 1.4-4.0, very thin- to very thick­ thickenings); 2) rays (of two distinct sizes walled, mostly with distinctly bordered pits or not, width, height and composition); 3) (fibre-tracheids), but infrequently with simple ground tissue fibres (pit border size and dis­ to minutely borderedpits (libriform fibres) in a tribution); 4) crystals (type, size, the types of few genera, the pit chambers 1-9 um in crystal-containing cells); 5) axial parenchyma diameter, in both radial and tangential walls, (amount). or mainly confined to the radial walls. Helical The Rosaceae are wood anatomically ra­ thickenings absent or present, elosely to wide­ ther diverse and there are few features com­ ly spaced, fine to coarse. mon to the whole family , as is apparent from Parenchyma scanty to abundant, frequent­ the following family description for the Chi­ ly apotracheally diffuse and scanty paratra­ nese representatives : cheal, but diffuse-in-aggregates or paren­ chyma in marginal or irregular zonate bands Growtb rings mostly distinct, rarely faint also present in some genera, in 3-5 (2-11)­ or indistinct, the boundaries wavy or not, celled strands. marked usually by rows of radially flattened Rays 5-17/mm, 1-16-seriate, of two latewood fibres, or also by differences in distinct sizes or of intergrading sizes. Uni­ vessel diameter and/or vessel frequency be­ seriate rays 2-16 cells high; multiseriate rays tween latewood and subsequent earlywood, 0.2-25.0 mm high, composed of square or or marginal parenchyma bands. Wood most­ square to procumbent body cells and upright ly diffuse-porous (rarely ranging to semi­ marginal cells, usually also with weakly dif­ ring-porous), infrequently ring-porous. Ves­ ferentiated sheath cells; or composed of pro­ sels 40-250 (8-520)/sq.mm, 5-99% cumbent body cells and 1-4(-7) rows of solitary, remainder in oblique , tangential and square to upright marginal cells; or composed radial multiples of 2-4(-12) vessels, round of procumbent cells only, sometimes with to oval and angular, tangential diameter 20­ one row of square marginal cells. Perforated 70 (16-123) um, radial diameter 30-80 ray cells almost always absent, but present in (20-160) um, walls mostly 1-2(-4) um one species. thick. Vessel element length 250-700 (200­ Crystals absent or sparse to abundant, 930) um, L/D ratio 4-30. Perforations al­ mainly prismatic, more rarely druses, sparse most exelusively simple in oblique to almost to abundant , medium-sized to large, in cham­ horizontal end walls, but sporadie scalariform bered and enlarged axial parenchyma cells and irregular multiple perforations present in and/or non-ehambered and non-enlarged ray some species. Intervessel pits mostly nonves- cells, usually one crystal per ray cell or cham-

Downloaded from Brill.com10/09/2021 05:06:33PM via free access 44 IAWA Bulletin n.s., Vol. 13 (1),1992 ber, chambered crystals in chains of 2-23. Rays 10 (7-12)/mm, more or less of two Silica bodies not observed. sizes: uniseriate rays .and (2-)3-4(-5)-seri­ Pith flecks sometimes present, usually dif­ ate rays. Uniserlate rays 5-6 (1-14) cells fuse or in tangential bands. high , composed of square to sometimes Traumatte ducts occasionally present, in weakly procumbent and upright marginal tangential bands, sometimes diffuse, mainly cells; multiserlate rays 390 (150-750) 11m present in earlywood or in the intermediate high, composed of procumbent body cells wood. and 1-4 rows of square to upright marginal cells (mainly heterogeneous Ill, occasionally heterogeneous ll) . Generic wood anatomical de8criptions Crystals common, prismatic, and medium­ sized, in non-enlarged procumbent and square Subfamily I: Spiraeoideae Agardh ray cells. Pith flecks and traumatic gum ducts absent. Exochorda Lindl. Note: The systematic position of the genus Material studied: E. giraldii Hesse: locality is still in dispute although traditionally it has unknown, CAFw 563. been placed in the Spiraeoideae. Wood ana­ Deciduous shrubs from East to North Chi­ tomically Exochorda is aberrant in the Spir­ na, often cultivated as ornamentals. aeoideae. The genus is more similar to the Growth rings distinct, marked by differ­ Prunoideae than to the Spiraeoideae. Like in ences in vessel diameter between latewood many Prunoideae, the rays in the genus are of and subsequent earlywood and rows of two sizes, lower than 1.0 mm and composed weakly flattened latewood fibres. Wood ring­ of procumbent body cells and 1-4 rows porous to semi-ring-porous, with one row of square to upright marginal cells, the pits in of large earlywood vessels. Latewood ves­ fibre-tracheids are mainly confined to the sels 185/sq.mm, 83% solitary, remainder radial walls, and helical thickenings occur in in oblique, radial and tangential multiples of vessel elements. However, the percentage of 2-3, round to oval or weakly angular, tan­ solitary vessels in the genus is higher than in gential diameter of earlywood vessels 41 the Prunoideae. (25-60) J.UD, radial diameter 48 (32-70) J.UD; tangential diameter of latewood vessels .23 Sorbaria (Ser.) A. Br. ex Aschers . (Fig. 8; (15- 34) J.UD, radial diameter 30 (18- 44) J.UD, Table 2) walls I-211m thick. Vessel element length Material studied: S. arborea Schneid. var. 370 (290-520) J.UD. Perforations exclusively subtomentosaRehd.: Tibet, CAFw 19346.­ simple in oblique end walls. Intervessel pits S. kirilowi (Regel) Maxirn.: Zhongtiaoshan, nonvestured, alternate, round, oval to polyg­ ·Sanxi, HEFw 137. - Sosorbifolia (L.) A. Br.: onal, 5-8 J.UD in diameter, with slit-like aper­ Shenyang, Liaoning, Qian Hong s.n., diam. tures. Vessel-ray and vessel-parenchyma sim­ 12mm. ilar to intervessel pits but half-bordered and Deciduous shrubs from Northeast, North slightly smaller. Helical thickenings well de­ and Southwest China, often cultivated as or­ veloped and closely spaced, mainly confined namentals. to narrow vessel elements. Gummy contents Growth rings distinct, marked by 1-4 absent. rows of radially flattened latewood fibres and Fibre-tracheids 740 (410-1030) J.UD long, locally inflated broad rays. Wood diffuse­ FIV ratio 2.0, very thick-walled, with dis­ porous to semi-ring-porous, Vessels 32-831 tinctly bordered pits of 3-5 11m in diameter, sq.mm, 77-85% solitary, remainder mainly mainly confined to radial walls. Helical thick­ in oblique and tangential multiples of 2-3, enings very fine, occasionally present in some round to oval, tangential diameter 45-73 fibre-tracheids. (16-110) 11m, radial diameter 54-89 (24­ Parenchyma common , apotracheally dif­ 135) J.UD, walls 1-3 J.UD thick. Vessel element fuse, diffuse-in-aggregates and scanty para­ length 370-410 (240- 520) um, LID ratio tracheal, in 3 (2- 6)-celled strands. 6 - 8. Perforations exclusively simple in

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Table 2. Variation in selected wood anatomical features of Sorbaria

2 3 4 5 6 7 8 9

S. arborea 32 73 410 3-7 1()()() 2.4 13 0.8 S. kirilowi 60 47 370 3-5 710 1.9 9 0.4 S. sorbifolia 83 45 370 + 2-4 660 1.8 6 0.6 1: Vessel frequency (lsq.mrn); 2: Average tangential vessel diameter (um); 3: Average vessel element length (um); 4: Multiple perforations present (+) or absent (-); 5: Size of intervessel pits (um); 6: Average fibre-tracheid length (um); 7: F/V ratio; 8: Width of the widest rays (number of cells); 9: Average multiseriate ray height (mrn). oblique to almost horizontal end walls, but in Notes: 1. Wood anatomically Sorbaria is S. sorbifolia sporadic irregular multiple per­ somewhat different from Spiraea and Stepha­ forations also found. Intervessel pits nonves­ nandra ofthe Spiraeoideae although it shares tured , altemate, round to oval, 3-7 um in a number of features comrnon to the two gen­ diameter, with slit-like, occasionally coales­ era. Sorbaria has wider vessel elements, a cent apertures . Vessel-ray and vessel-paren­ significantly lower L/D ratio, a lower vessel chyma pits similar to intervessel pits but half­ frequency, more abundant axial parenchyma, bordered and slightly smaller. Helical thick­ and lower multiseriate rays than Spiraea and enings absent. Gumrny contents present in Stephanandra. Moreover, rays in Sorbaria are some vessels ofS. aboreavar. subtomentosa composed of procumbent cells only, or rarely and S. kirilowi . have one row of square to upright marginal Fibre-tracheids 660-1000 (410-1280) um cells. However, Suzuki et al. (1991) reported long, F/V ratio 1.8-2.4, medium thick­ more strongly heterogeneous rays in Sorbaria walled, with distinctly bordered pits of 3-5 tomentosa from the Himalayas. um in diameter, in radial (sometimes densely 2. Scattered multiple perforations in some placed) and tangential walls. Helical thicken­ species of Sorbaria were recorded by Met­ ings absent. calfe and Chalk (1950). However, none were Parenchyma abundant, predominantly apo­ observed in the Chinese species. tracheally diffuse , but diffuse-in-aggregates 3. Differences in some anatomical features and scanty paratracheal also present, in 3-4 (vessel frequency, vessel diameter, vessel ele­ (2-6)-celled strands . ment length and fibre length) between S. ar­ Rays 5-10 (1-13)/mm, of two distinct borea var. subtomentosa from Tibet and the sizes: narrow and low rays 1-2(-3)-seriate, remaining two species are very appreciable and wide and tall rays (4-)5-13-seriate (the (see Table 2). widest rays 6-seriate in S. sorbifolia to 13­ seriate in S. arborea var. subtomentosa). Uni­ Spiraea L. (Figs. 10, 25, 26, 40-42, 45, 52, seriate rays 3-4 (1-13) cells high, composed 57; Tables 3-5) of upright and square cells; multiseriate rays Material studied : S. blumei G. Don: Hu­ 350-790 (70-2100) um high, composed of angshan, Anhui, Huang Chenlin s.n., diam. procumbent cells only, or rarely with one 15 mrn, alt. 500 m. - S. cantoniensis Lour. : row of square to upright marginal cells (main­ Hefei , Anhui, S.Y. Zhang s.n ., cult., diam. ly homogeneous, occasionally heterogeneous 18 mrn, alt. 50 rn. - S. chamaedryfolia L.: III). Changbaishan, Jining, Qian Hong s.n., 10 Crystals absent. Pith flecks rare in S. ar­ mrn diam. - S. chinensis Maxirn.: Huoshan , borea var. subtomentosa and S. kirilowi, Anhui, He Yunhe s.n. , diam . 15 mm, alt. small, in tangential bands or diffuse. Pith 200 m. - S. japonica var. pinnatifida T.T. Yu flecks not observed in S. sorbifolia. Traum­ & L.T. Lu: Qingliangfeng, Anhui, Huang atic gum ducts .absent. Chenglin s.n., diam. 10 mrn, alt. 400 m. - S.

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Tab1e 3. Variation in se1ected wood anatomical features of Spiraea.

1 2 3 4 5 6 7 8 S. blumei 73 99 31 300 6 1.1 R S. cantoniensis 90 99 19 290 7 2.0 R S. chamaedryfolia 198 71 22 260 + 6 4.4 RA S. chinensis 62 91 43 7 2.6 RA S.japonica 135 82 21 350 7 2.7 R var. pinnatifida S. prunifolia 104 92 30 300 6 4.7 R var. simplicijlora 65 86 25 290 11 2.4 R S. salicifolia 385 78 24 290 + 6 1.7 S. thunbergii Liu Xiumei s.n. 43 97 29 390 7 4.2 Qian Hong s.n . 136 69 24 260 6 3.3 R S. trilobata 53 98 28 280 6 1.7 R 1: Vesse1frequency (/sq.mm); 2: Percentage of solitary vesse1s; 3: Average tangential vesse1 diameter (1JlIl); 4: Average vesse1element 1ength(1JlIl); 5: Scalariform and/or irregular perfora- tions present (+) or absent (-) ; 6: Width ofthe widest rays (number of cells); 7: Average mu1ti- seriate ray height (mm); 8: Prismatic crystals present in ray cells (R), axial parenchyrna (A), in both (RA) or absent (-). prunifolia Sieb. & Zucc.: Tunxi, Anhui, Ye or weakly angular, tangential diameter 19-43 Shuyou s.n., diam. 9 mm, alt. 200 m. - S. (10-60) 1JlIl, radial diameter 26-53 (15-75) prunifolia Sieb. & Zucc. var. simplicijlora um, walls 1-3 um thick. Vesse1 element Nakai: Hefei, Anhui, S.Y. Zhang s.n., cult., length 260-390 (160-720) um. L/D ratio diam. 22 mm, alt. 50 m. - S. salicifolia L.: 8-17. Perforations exc1usive1y simple in Changbaishan, Jining, Qian Hong s.n., 13 oblique to almost horizontal end walls, but in mm diam. - S. thunbergii Sieb. ex Blume: S. chamaedryfolia and S. salicifolia sporadie Huangshan, Anhui, Liu Xiumei s.n., diam. scalariform and irregular multiple perfora­ 18 mm, alt. 500 m; Shenyang, Liaoning, Qi­ tions with 1-6 bars also noted mainly in sec­ an Hong s.n., diam. 10 mm. - S. trilobata L.: ondary xylem nearby the pith. Intervessel pits Tunxi, Anhui, Ye Shuyou s.n., diam. 18 mm, nonvestured, or vestured in S. blumei, S. alt. 200 m. chinensis, S. japonica var. pinnatifida, S. Deciduous shrubs from Northeast, North­ prunifolia var. simp/icijlora, S. salicifolia, west, Southwest and East China, usually cul­ and S. thunbergii , vestures mainly confined tivated as ornamentals. to rare pits nearby the perforation rims, pits Growth rings distinct, occasionally with alternate, oval to polygonal, 3-5 um in wavy boundaries, marked by one to several diameter, with slit-like apertures, apenures rows of weakly radially flattened latewood occasionally coalescent in S. chamaedryfolia, fibres, differences in vessel diameter between S. prunifolia var. simplicijlora and S. thun­ latewood and subsequent earlywood, and in bergii. Vesse1-ray and vessel-parenchyma some species by locally inflated broad rays pits simi1ar to intervessel pits but half-bor­ as weIl. Wood semi-ring-porous to diffuse­ dered. Helical thickenings absent. Gummy porous. Vessels sometimes in a weakly pro­ contents rarely present nounced radial to oblique pattern in a few Ground tissue composed of two types of species, 43-385 (mostly 50-136)/sq.mm, fibres: fibre-tracheids and libriform fibres 69-99% (mostly above 90%) solitary, re­ (for details, see note below), 400-590 (230­ mainder in oblique and tangential, occasion­ 770) um long, F/V ratio 1.5-1.9, medium ally in radial multiples of 2-3, round, oval thick- to very thick-walied, with simple to

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Table 4. Comparison of quantitative features offibre-tracheids and librifonn fibres of Spiraea.

fibre-tracheids librifonn fibres 2 3 4 5 6 7 2 3 4 5 6 7

S. blumei 590 13.5 1.9 5.8 2.0 0.33 3 581 14.9 4.3 5.3 1.9 0.80 1- 2 S. cantoniensis 514 13.2 2.6 5.3 1.8 0.49 3 496 14.6 6.4 4.1 1.7 1.56 1-2 S. chamaedryfolia 416 10.0 2.4 3.8 1.6 0.63 3 471 12.3 4.4 3.9 1.8 1.13 1- 2 S. japo nica var. pinnatifida 446 13.1 5.0 4.1 1.3 1.22 3 553 18.3 7.9 5.2 1.6 1.52 1- 2 S. prunifolia 478 12.6 2.7 5.0 1.6 0.54 3 462 14.5 4.6 5.0 1.5 1.92 1-2 var. simpliciflora 408 10.6 2.6 4.0 1.4 0.65 3 499 13.3 4.7 4.3 1.7 1.09 1-2 S. salicifolia 448 13.0 2.9 5.0 1.6 0.58 3-4 516 14.3 3.9 5.2 1.8 0.75 1- 2 S. thunbergii Liu Xiumei s. n, 569 13.7 2.8 5.4 1.5 0.52 3 556 15.0 5.9 4.6 1.4 1.28 1-2 Qian Hong s.n. 357 11.8 2.9 4.5 1.4 0.64 3 413 13.0 4.3 4.4 1.6 0.98 1-2 S. trilobata 436 13.0 2.0 5.5 1.6 0.36 3 499 15.9 6.0 4.9 1.8 1.22 1- 2

1: Length (Jl.III); 2: Diameter (JUI1); 3: Lumen diameter UJ,m); 4: Wall thickness (JUI1); 5: F/V ratio ; 6: Lumen diameter/wall thickness ratio (L/ W ratio); 7; Pit size (JUI1).

Table 5. Analysis of variance of quantitative Rays 9-15 (5-19)/mm, of two distinct features of librifonn fibres and fibre-tracheids sizes: narrow and low rays 1(-2)-seriate, in Spiraea. and wide and tall rays 4-6(-1l)-seriate (the Parameter Degrees of Mean F-value widest rays 6- to 7-seriate in all species ex­ freedom square cept in S. prunifoliavar. simpliciflora where the widest rays are up to l l-seriate). Uniseri­ Length 1 7372.8 1.9 ate rays 3-8 (1-21) cells high, composed 18 3901.3 exclusively of upright cells; multiseriate rays Diameter 1 23.3 10.7** 1.1-4.7 (0.3-12.5) mm high, composed of 18 2.2 square to procumbent body ray cells, rows of Lumen diameter 1 32.8 26.1** upright marginal cells and weakly differen­ 18 1.3 tiated sheath cells. Perforated ray cells noted Wall thickness I 0.04 0.09 in S. chamaedryfolia. 18 0.4558 Crystals sparse to abundant, prismatic, usu­ ally medium-sized, occasionally elongated, F/V ratio I 0.05 1.52 mostly present in non-eharnbered ray cells, 18 0.033 one crystal per ray cell, but prismatic and L/W ratio I 1.51 21.9 ** fragmented crystals noted in chambered, non­ 18 1.25 enlarged axial parenchyma cells only in S. ** = difference highly significant. chamaedryfo lia, 1-5 crystals per chamber, crystals not observed in S. saLicifolia and S. thunbergii (Liu Xiumen s.n.). Pith flecks distinctly bordered pits of 1-4 11m in diam­ present in S. chamaedryfolia and S. salici­ eter, in radial (densely placed) and tangen­ folia , small to large, usually in tangential tial walls. Helical thickenings absent. Sparse bands. Traumatic gum ducts absent septate librifonn fibres noted in S. chinensis, Notes: 1. The occurrence of librifonn fi­ S. chamaedrifolia, S. japonica var. pinnati­ bres in Spiraea was also reported by Solereder fida and S. prunifolia var. simplicijlora. (1908), Metealfe and Chalk (1950), Carlquist Parenchyma usually scanty, apotracheally (1988a), and Schweingruber (1990). The diffuse and scanty paratracheaI, in 2-3 (5)­ ground tissue in all Chinese species studied is celled strands. composed of librifonn fibres with sparse,

Downloaded from Brill.com10/09/2021 05:06:33PM via free access 48 IAWA Bulletin n.s ., Vol. 13 (1),1992 simple to minutely bordered pits of 1-2 um 71) um, walls I-211m thick. Vesse1 element in diameter, main1y confined to radial walls, 1ength 540-630 (220-980) 11m. L/D ratio and fibre-tracheids with densely placed dis­ 16-18. Perforations almost exclusively sim­ tinctly bordered pits of 3(-4) um in diameter, ple in oblique to almost horizontal end walls, numerous in both radial and tangential walls. but sporadic scalariform perforations with The libriform fibres are mainly present in less than 5 bars also noted in one sarnple earlywood and intermediate wood. The fibre­ (Huang Chenglin s.n.). Intervessel pits non­ tracheids are mainly present in the latewood. vestured, alternate, round , oval to polygonal, As shown in Tables 4 and 5, there are no sig­ 3-6 11m in diameter, with slit-1ike apertures, nificant differences in cell length between apertures sometimes coa1escent. Vessel-ray libriform fibres and fibre-tracheids. But the and vessel-parenchyma similar to intervessel libriform fibres are significantly wider than pits but ha1f-bordered. Helical thickenings the fibre-tracheids. There are no significant absent. Gumrny contents absent. differences in cell wall thickness between the Fibre-tracheids 850-910 (600-1340) 11m two types of elements. However, the lumen long, F/V ratio 1.3-1.7, medium thick- to diameter of the libriform fibres is larger than very thick-walled, with distinctly bordered that ofthe fibre-tracheids, Furthermore, lumen pits of 4-7 11m in diameter, densely placed in diameter/wall thickness ratio (L/W ratio) in radial and tangential walls. Helical thickenings the libriform fibres is significantly larger than absent . in the fibre-tracheids (Tables 4 & 5). In other Parenchyma rather sparse (Huang Cheng­ words, the fibre-tracheids are mostly medium lin s.n.) to abundant (He Yunhe s.n.), main­ thick-walled, whereas the libriform fibres are ly apotracheally diffuse, but scanty paratra­ mostly thin-walled according to the classifi­ cheal and diffuse-in-aggregates also present cation by Van Vliet (1976). in one sample (He Yunhe s.n.), in 3-4 (2-8)­ 2. Metealte and Chalk (1950) stated that celled strands. Spiraea includes some ring-porous or semi­ Rays 15-17 (12-20)/mm, of two dis­ ring-porous species. However, all samples tinct sizes: low and narrow rays 1(-2)-seriate, observed here are semi-ring-porous to diffuse­ and wide and tall rays 3-6-seriate. Uniseri­ porous. Diffuse-porous to slightly semi-ring­ ate rays 10-15 (2-36) cells high, composed porous wood in S. salicifolia and S. hyperi­ of upright cells; multiseriate rays 2.6-4.1 cifolia was also reported by Schweingruber (0.4-15.0) mm high , composed of square (1990) . Diffuse-porous rootwood of the and upright cells, without procumbent cells. genus was described by Cutler et al. (1987). Crystals sparse to common, prismatic, oc­ 3. The report of vestured pits in Spiraea is casionally elongated, medium-sized, present new for Rosaceae . in non-enlarged square ray cells. Pith flecks and traumatic gum ducts absent. Stephanandra Sieb. & Zucc, Notes: 1. Helical thickenings in ground tis­ Material studied: S. chinensis Hance: Huo­ sue fibres in the genus were recorded by Met­ shan, Anhui, He Yunhe s.n ., diam. 15 mm, calfe and Chalk (1950) , but we did not ob­ alt. 100 m; Huangshan, Anhui, Huang Cheng­ serve such thickenings in the Chinese species. lin s.n., diam. 12 mm, alt. 700 m. 2. The difference in the amount ofaxial Deciduous shrubs from South to North­ parenchyma between the two sampIes is ap­ east China, sometimes cultivated as ornarnen­ preciable. .tals. Growth rings faint to distinct, with more Subfamily ll: Maloideae Weber or less wavy boundaries, marked by rows of weakly radially flattened latewood fibres. Amelanchier Medic. Vessels diffuse, 109-125/sq.mm, 67-77% Material studied: A.sinica (Schneid.)Chun: solitary, remainder mainly in oblique and tan­ Shanxi, CAFw 18850. gential multiples of 2-4, oval to round or Deciduous shrubs or trees from East, weakly angular, tangential diameter 31-40 Northwest and Central China, often culti­ (16-53) 11m, radial diameter 40-51 (21- vated as ornamentals.

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Growth rings distinct, marked by 1-4 marginal cells (homogeneous and heterogen­ rows of radially flattened latewood fibres . eous Ill). Vessels diffuse, 290/sq.mm, 79% solitary, Crystals absent, Pith flecks present,Trau­ remainder mainly in oblique multiples of matic gum duets absent, 2-3, round or oval to angular, tangential diameter 27 (13-35) um, radial diameter 31 Chaenomeles Lind!. (Fig. 22; Table 6) (18-46) um, walls I-211m. Vessel element Material studied:C. sinensis (Thouin) length 630 (520-780) um, LID ratio 23. Koehne: Hefei, Anhui, S.Y. Zhang s.n., Perforations almost exclusively simple in cult., diarn. 20 mm, alt. 50 m. - C. speciosa obl ique end walls, but sporadie irregular (Sweet) Nakai: Hefei, Anhui, S.Y. Zhang multiple perforations also noted. Intervessel s.n., cult., diam. 11 mm, alt 50 m. - C. thi­ pits nonvestured, altemate, round to oval, betica Yu: Tibet, CAFw 19332. 3-5 11m in diameter, with slit-like apertures, Deciduous or semi-evergreen shrubs or the apertures occasionally coalescent, Vessel­ small trees mainly from Northwest and South­ ray and vessel-parenchyma pits similar to west China, widely cultivated as ornamentals intervessel pits but half-bordered. Helical and fruit trees. thickenings fine and irregularly spaced, Growth rings distinct to faint, marked by throughout body of vessel elements, often 1-4 rows of weakly radially flattened late­ associated with pit apertures. Gummy con­ wood fibres. Vessels diffuse, (36-)149­ tents absent . 264/sq.mm, 85-97% solitary, remainder in Fibre-tracheids 860 (600-1220) um long, oblique, tangential and radial multiples of F/V ratio 1.4, medium thick- to very thick­ 2-3, round or oval to angular, tangential walled, with distinctly bordered pits of 5-6 diameter 22-63 (12-107) um, radial diam­ umin diameter, in radial and tangential walls. eter 28-80 (16-143) um, walls 1-3 um Helical thickenings absent, thick. Vessel element length 350-550 (220­ Parenchyma moderately abundant, apo­ 900) um, LID ratio (9-)15-19. Perforations tracheally diffuse, diffuse-in-aggregates, and exc1usively simple in oblique to almost hori­ scanty paratracheal, in 5 (3- 1O)-celled zontal (only in C. thibetica) end walls, but strands. sporadie scalariform perforations also noted Rays 8-9 (7-ll)/mm, 1-2-seriate. Uni­ in C. speciosa. Intervessel pits nonvestured, seriate rays 4 (1-11) cells high, composed of alternate, round to oval , 5-9 um in diameter, procumbent cells, occasionally with one row with slit-Iikeapertures. Vessel-ray and ves­ of square marginal cells; biseriate rays 180 sel-parenchyma pits similar to intervessel (70 - 430) 11m high, composed of procum­ pits but half-bordered and slightly smaller. bent cells only, or with one row of square Helical thickenings well developed and usu-

Table 6. Variation in selected wood anatomical features ofChaenomeles.

2 3 4 5 6 7 8 9 10 11 12 C. sinensis 149 22 430 + 910 2.1 4-6 + 0.2 11 & rn - C. speciosa 264 23 350 + + 670 1.9 3-4 + 0.2 11 & III + C. thibetica 36 63 550 920 1.7 4-5 0.4 Ito III +

1: Vessel frequency (lsq.mm); 2: Average tangential vessel diameter (um); 3: Average vessel length (um); 4: Multiple perforations present (+) or absent (-); 5: Helical vessel wall thickenings present (+) or absent (-); 6: Average fibre-tracheid length (um) ; 7: F/V ratio; 8: Size of pits in fibre-tracheids (um): 9: Helica1 thickenings present in fibre-tracheids (+) or absent (-); 10: Aver­ age multiseriate ray height (mm); 11: Ray types; 12: Crystals present (+) or absent (- ).

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ally closely spaced, throughout body of ves­ 271-315/sq.mm, 69-86% solitary, remain­ sei elements in C. sinensis and C. speciosa, der mainly in oblique and tangential multiples thickenings not observed in C. thibetica. of 2-3, oval, round or angular, tangential Gummy contents absent, diameter 24-25 (13-40) J.LIll, radial diameter Fibre-tracheids 670-920 (470-1260) J.LIll 31-33 (15-53) um, walls 1-2 um thick. long, F/V ratio 1.7-2.1, medium thick- to Vessel element length 300-420 (180-600) very thick-walled, with distinctly bordered um, LID ratio 12-17. Perforations exclu­ pits of 3-6 J.LIll in diameter, in radial and tan­ sively simple in oblique end walls. Interves­ gential walls. Helical thickenings fine 10 con­ sei pits nonvestured, alternate, round, 3-6 spicuous, present in some to almost all fibre­ J.LIll in diameter, with slit-like apertures. Ves­ tracheids of C. sinensis and C. speciosa. sel-ray and vessel-parenchyma pits similar to Parenchyma abundant, apotracheally dif­ intervessel pits but slightly smaller and half­ fuse, diffuse-in-aggregates and scanty para­ bordered or with slightly reduced borders. tracheal, in 4-5 (2-1O)-celled strands. Helical thickeningsweIl developed and close­ Rays 10-15 (7-17)/mm, 1-3-seriate. Iy spaced, throughout body of vessel ele­ Uniseriate rays 4-6 (1-13) cells high, com­ ments. Gummy contents present in a few posed of square and upright ceIls; 2-3-seri­ vessels of C. microphyllus. ate rays 200-390 (70-900) um high, com­ Fibre-tracheids 580-820 (410-1000) um posed ofprocumbent body cells and 1-2(-6) long, F/V ratio 1.9, medium thick- to very rows of square to upright marginalcell (main­ thick-walled, with distinctly bordered pits of ly heterogeneous Ill, rarely heterogeneous n 4-5 J.LIll in diameter, in radial and tangential in C. sinensis and C. speciosa, but heterogen­ walls. Helical thickenings present in almost eous rn to I are all common in C. thibetica). all fibre-tracheids, Crystals abundant in C. speciosa and C. Parenchyma abundant, apotracheally dif­ thibetica, prismatic, medium-sized to large, fuse, diffuse-in-aggregates and scanty para­ in chambered axial parenchyma ceIls, one tracheal, in 3-4 (2-7)-celled strands. crystal in each enlarged, sclerified chamber; Rays 12 (9-15)/mm, 1-3-seriate. Uni­ chains of up to 23 chambers, crystals absent seriate rays 3-5 (1-17) cells high, composed in C. sinensis. Pith flecks sparse in C. sinen­ of square (to weakly procumbent) cells and sis and C. thibetica, but absent in C. speciosa. upright ceIls; 2-3-seriate rays 180-190 Traumatic gum ducts absent. (50-450) um high, composed of procum­ Note: As shown in Table 6, C. thibetica bent body cells and 1-3 rows of square (to from Tibet has much larger vessel elements, upright) marginal cells (mainly heterogeneous and a much lower vessel frequency than the Ill, rarely heterogeneous Il), remaining two species. Chaenomeles thibeti­ Crystals absent in C. microphyllus; sparse ca also stands out in absence of helical thick­ in C. multiflora, prismatic, medium-sized to enings and ray composition. The latter differ­ large, in chambered axial parenchyma ceIls, ences are probably related to differences in one crystal in each enlarged and sclerified sampIe size (mature stemwood in C. thibeti­ chamber; chains of up to 11 chambers. Pith ca; thin stern in other two species). flecks absent in C. multiflorus, but common in C. microphyllus. Traumatic gum ducts ab­ Cotoneaster B. Ehrhart sent. Material studied: C. microphyllus Wall. ex Notes: 1.The drawings by Greguss (1959) Lindl.: Tibet, CAFw 19327 diam. 30 mm- indicate occurrence of libriform fibres in Co­ C. multiflorus Bunge: Sichuan, CAFw 561. toneaster p.p. No libriform fibres, however, Deciduous, evergreen or semi-evergreen were observed in the sampies we exarnined. shrubs, sometimes small trees, mainly from 2. Semi-ring-porous wood in C. nummu­ western regions and Southwest China. laria was reported by Fahn et al. (1986). A Growth rings distinct, marked by 1-4 study of several species by Schweingruber rows of radially flattened latewood fibres. (1990) indicated that porosity in the genus Wood diffuse-porous in C. microphyllus to also ranges from diffuse- to semi-ring-por­ semi-ring-porous in C. multiflorus. Vessels ous .

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Table 7. Variation in selected wood anatomical features of Crataegus .

2 3 4 5 6 7 8 9 10

C. aurantia 304 71 27 390 780 2.0 + 2 C. cuneata 197 74 32 370 650 1.7 + 3 + C. hupehensis 211 80 28 410 + 750 1.8 + 3 + C. maximowiczii 520 65 22 350 + 730 2.0 + 2 C.oresbia 456 44 27 600 860 1.4 + 2 + C. pinnatifida 268 60 35 380 + 750 2.0 3 C. shensiensis 222 76 29 420 + 820 2.0 + 3(-4) + C. wattina 140 83 33 310 800 2.6 + 3(-4) C. wilsonii 184 76 31 390 780 2.0 + 3(-4) +

1: Vessel frequency (/sq.mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (11m); 4: Average vessel element length (11m); 5: Multiple perforations present (+) or absent (-); 6: Average fibre-tracheid length (11m); 7: F/V ratio; 8: Helical thickenings present in fibre-tracheids (+) or absent (-); 9: Width of the widest rays (number of cells); 10: Crystals present (+) or absent (-).

3. Vessel frequency and ray frequency in round to angular, tangential diameter 22-35 five species studied by Greguss (1959) are (13-58) 11m, radial diameter 27-48 (15­ significantly higher than in our material . 72) 11m, walls I-211m thick . Vessel element length 310-600 (110 -830) 11m. L/D ratio Crataegus L. (Figs. 38,49, 50; Table 7) (9-)11-22. Perforations exclusively simple Material studied: C. aurantia Pojark.: in oblique to almost horizontal end walls ex­ Zhongtiaoshan, Shanxi, HEFw 116. - C. cept in C. hupehensis,C. maximowiczii, C. cuneata Sieb.: Huoshan, Anhui, He Yunhe pinnatifida and C. shensiensis where sporadic s. n., diam. 15 mm, alt. 200 m. - C. hupe­ scalariform and/or irregular multiple perfora­ hensis Sarg.: Huoshan, Anhui, He Yunhe tions with 2-5 bars were also found. Inter­ s. n., diam. 20 mm, alt. 200 m. - C. maxi­ vessel pits nonvestured, usually alternate, mowiczii Schneid.: Changbaishan, Jining, mostly round, 6-9 11m in diarneter, with slit­ Qian Hong s.n., diam. 13 mm. - C. oresbia like apertures. Vessel-ray and vessel-paren­ W.W. Smith: Yunnan, CAFw 3239. - C. chyma pits half-bordered and slightly smaller, pinnatifida Bunge: Northeast China, North­ 4-6 11m in diameter. Helical thickenings fine east Forestry University s.n. - C. shensien­ and closely spaced, usually confined to tails sis Pojark.: Zhongtiaoshan, Shanxi, HEFw ofsome vessel elements; weil developed heli­ 117. - C. wattina Hemsl, & Lace: Zhongtiao­ cal thickenings throughout body of vessel shan, Shanxi, HEFw 118. - C. wilsonii elements in C. aurantia and C. oresbia, heli­ Sarg.: Zhongtiaoshan, Shanxi, HEFw 119. cal thickenings not observed in C. pinnatifi­ Deciduous, rarely serni-evergreen shrubs da.Gummy contents present in a few vessels or small trees mainly from North and South­ of C. hupehensis and C. wilsonii. west China. Fibre-tracheids 650-860 (490-1290) 11m Growth rings distinct, marked by rows of long, F/V ratio 1.4-2.0(-2.6), very thin - to radially flattened latewood fibres. Vessels dif­ very thick-walled, with distinctly bordered fuse , 140-520/sq.mm (mostly 140-300/ pits of 4-6 11m in diameter, in radial and tan­ sq.mm), 44-83% (all above 60% except in gential walls. Helical thickenings fine to con­ C. oresbia) solitary, remainder in oblique, spicuous, present in all species except in C. radial and tangential multiples of 2-3, oval , pinnatifida.

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Parenchyma usually abundant (except in J.1m, walls 1-2 um thick. Vessel element C. oresbia with scanty parenchyma), mainly length 760 (360-720) um, L/D ratio 29. Per­ apotracheally diffuse, but diffuse-in-aggre­ forations exclusively simple in oblique end gates and scanty paratraehea1 parenchyma also walls . Intervessel pits nonvestured, alternate, present, in 3-5 (1-8)-celled strands. round to oval, 5-6 um in diameter, with slit­ Rays 10-15 (7-17)/mm, 1-3(-4)-seri­ like apertures, Vessel-ray and vessel-paren­ ate. Uniseriate rays 4-13 (1-41) cells high, chyma pits similar to intervessei pits but half­ composed of procumbent cells only, or oc­ bordered and slightly smaller. Helical thick­ casionally with one row of marginal cells; 2-3 enings weakly developed, sometimes irregu ­ (-4)-seriate rays 190-300 (50-680) J.1m lar, present in part of vessel elements. Gum­ high, composed of procumbent cells only, or my contents present in a few vessels. rarely with one row of square marginal cells Fibre-tracheids 1060 (640-1360) J.1m (mainly homogeneous, occasionally hetero­ long, F/V ratio 1.4, very thick-walled, with geneous III, but in branchwood of C. cune­ distinctly bordered pits of 4-5 J.1m in diam­ ata heterogeneous 11 also present) . eter, in radial and tangential walls. Helical Crystals present in most species, usual­ thickenings present in most fibre-tracheids. ly sparse, prismatic, occasionally elongated, Parenchyma abundant, apotracheally dif­ large, present in chambered axial parenchyma fuse, diffuse-in-aggregates and scanty para­ cells or idioblasts, one crystal in each enlarg­ tracheal, in 5 (3-9)-celled strands. ed, sclerified chamber; chains of up to 13 Rays 10 (8-14)/mm, 1-2-seriate. Uni­ chambers, crystals not observed in C. auran­ seriate rays 5 (1-18) cells high, composed of tia, C. maximowiczii, C. pinnatifida and C. procumbent cells and with one row of square wattina. Pith flecks common, but absent in marginal cells; biseriate rays 250 (90-450) C. cuneata, C. hupehensis and C. oresbia. JlID high, composed of procumbent cells Traumatic gum ducts absent. only, or with one row of square marginal Notes: 1. The drawings by Greguss (1959) cells (predominantly heterogeneous m, rarely indicate absence of helical thickenings in fibres homogeneous). of Crataegus p.p. However, we observed Crystals abundant, prismatic, large, pres­ them in almost all Chinese samples studied. ent in chambered axial parenchyma cells, one In addition, the drawings show incidence of crystal in each enlarged, sclerified chamber; libriform fibres in the genus, but they were chains of up to 20 chambers. Pith flecks and not noted in OUT samples, traumatic gum ducts absent. 2. Both prismatic crystals and druses in Notes: 1. The drawings by Greguss (1959) the genus were recorded by Schweingruber show occurrence of libriform fibres in Cydo­ (1990). However, druses were not observed nia species but absence ofhelica1 thickenings in the Chinese specimens. in fibres, which differs from our observations. 3. Intervessel pits opposite as reported by 2. Sporadic multiple perforations were re­ Jacquiot et al. (1973) in C. oxyacantha and corded by Metealfe and Chalk (1950) and C. monogyna do not occur in the Chinese Fahn et al. (1986) in the genus . However, species. multiple perforations were not observed in the Chinese species. Cydonia Mill. Material studied: C. oblonga Mill.: Honan, Dichotomanthes Kurz (Figs. 51, 55) CAFw 5653. Material studied: D. tristaniaecarpa Kurz: Monotypic genus, deciduous shrubs or Yunche, Yunnan, Yunnan Academy of For­ small trees, cultivated in temperate regions . estry 346, alt. 1800. Growth rings distinct, marked by rows of Monotypic genus of evergreen shrubs to weakly radially flanened latewood fibres. small trees, native in Southwest China. Vessels diffuse, 134/sq.mm, 95% solitary, Growth rings faint, marked by rows of remainder mainly in oblique pairs, round to weakiy radially flanened latewood fibres. oval or weakly angular, tangential diameter Vessels diffuse, 187/sq.mm, 80% solitary, 26 (16-37) um, radial diameter 33 (23-50) remainder mainly in oblique, rarely tangential

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 53 and radial multiples of 2-3, angular, tangen­ Docynia Dcne. tial diameter 40 (18-45) 11m, radial diameter Material studied: D. indica(Wall.) Decne.: 43 (24-60) um, walls I-211m thick. Vessel Sichuan, CAFw 6570. element length 560 (380-780) um, L/D ratio Evergreen or semi-evergreen trees from 14. Perforations exclusively simple in ob­ Southwest China. lique to almost horizontal end walls. Inter­ Growth rings distinct, marked by rows of vessel pits nonvestured, alternate, round, 3-5 radially flattened latewood fibres. Vessels 11m in diameter, with slit-like apertures. Ves­ diffuse, 9O/sq.mm, 87% solitary, remainder sel-ray and vessel-parenchyrna pits similar in oblique and tangential pairs, round to oval to intervessel pits but half-bordered or with or weakly angular, tangential diameter 39 slightly reduced borders. Helical thickenings (20- 57) um, radial diameter 57 (28- 84) um, closely spaced or irregular, throughout body walls I-211m thick. Vessel element length of most vessel elements. Gummy contents 270 (170-380) 11m. L/D ratio 7. Perfora­ present in a few vessels. tions exclusively simple in oblique end walls. Fibre-tracheids 1100 (830-1500) 11m Intervessel pits nonvestured, alternate, round long, F/V ratio 1.9, medium thick-walled, to oval, 5-6 11m in diameter, with slit-like with distinctly bordered pits of 5-8 11m in apertures. Vessel-ray and vessel-parenchyma diameter, in radial and tangential walls. Heli­ pits similar to intervessel pits but half-border­ cal thickeningspresent in most fibre-tracheids. ed and slightly smaller. Helical thickenings Parenchyrna abundant, apotracheally dif­ fine and usually closely spaced, confined to a fuse, diffuse-in-aggregates and scanty para­ few narrow vessel elements. Gummy con­ tracheal, in 4 (2-7)-celled strands. tents absent. Rays 10 (7-14)/mm, 1-3-seriate. Uni­ Fibre-tracheids 620 (330-830) 11m long, seriate rays 3-4 (1-9) cells high, composed F/V ratio 2.3, medium thick-to very thick­ of square 10 weakly procurnbent cells and up ~ walled, with distinctly bordered pits of 5-6 right cells; 2-3-seriate rays 360 (70-600) um in diameter, in radial and tangential walls. um high, composed of procumbent body cells Helical thickenings absent. and 1-2(-4) rows of square (to upright) Parenchyma abundant, apotracheally dif­ marginal cells (heterogeneous mand Il), fuse, diffuse-in-aggregates and scanty para­ Crystals moderately abundant, prismatic, tracheal, in 3 (2-5)-celled strands. large, present in chambered axial parenchyma Rays 11 (8-13)/mm, 1-3-seriate. Uni­ cells or idioblasts, one crystal in each en­ seriate rays 3 (1-11) cells high, composed of larged and sclerified chamber; chains of up 10 procumbent cells; 2-3-seriate rays 260 (80­ 4 chambers. Pith flecks and traumatic gum 560) um high, composed of procurnbent cells ducts absent. only,or with one row of square marginal Notes: 1. This is the first wood anatomi­ cells (homogeneous and heterogeneous Im. cal description of Dichotomanthes. Crystals abundant, prismatic, large, in 2. Dichotomanthes has been considered by chambered axial parenchyrna cells, one crys­ different authors as belonging to the subfam­ tal in each enlarged, sclerifiedchamber; chains ilies Maloideae, Prunoideae or Spiraeoideae of up to lOchambers. Pith flecks occasional­ (Challice 1981). Gladkova (1969), on the ba­ ly present. Traumatic gurn duets absent. sis of morphological characters of fruits and pollen grains, even suggested a separate sub­ family for the genus. Yu (1974) put the genus Eriobotrya Lindl. (Fig. 30; Table 8) in the Maloideae, based on macromorpho­ Materialstudied: E. cavaleriei (Levl.)Rehd.: logical characters, Challice (1974) reported Guangdong, CAFw 14; Guangxi, South Chi­ that the occurrence of flavone C-glycosides na Agricultural University 844. - E. deflexa in the genus indicates strong affinity with the (Hemsl.) Nakai: Taiwan, CAFw 7306. - E. Maloideae and regarded it as a relic of the fragtans Champ.: Guangdong, CAFw 1154. primitive Maloideae. The wood structure of -E.japonica (Thunb.) Lindl.: Taiwan, CAFw the genus perfectly fits in the Maloideae with­ 7627; Guangxi , Guangxi Forestry Coll. s.n.; out indicating any specially primitive status. Fujian, CAFw 19781; Guangdong, FRIGw

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Table 8. Variation in selected wood anatomical features of Eriobotrya.

2 3 4 5 6 7 8 E. cavaleriei CAFw14 49 96 48 930 + 1570 1.7 0.3 844 48 98 57 910 1600 1.8 0.4 E. deflexa 160 73 38 770 + 980 1.3 0.3 Eifragrans 142 92 31 740 + 1200 1.6 0.3 E.japonica CAFw 7627 182 90 31 580 1050 1.8 0.3 Guangxi s.n. 171 96 30 640 + 1020 1.6 0.3 CAFw 19781 256 93 25 490 910 1.8 0.2 FRIGw 1060 256 90 26 640 1200 1.9 0.2 HEFw 293 178 89 22 480 1000 2.1 0.2 HEFw294 141 85 28 570 1190 2.1 0.3

1: Vessel frequency (lsq .mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (um); 4: Average vessel element length (um); 5: Multiple perforations present (+) or absent (-); 6: Average fibre-tracheidlength (um); 7: F/V ratio; 8: Average multiseriate ray height (mm).

1060; Anqing, Anhui, HEFw 293, alt. 200 throughout body of vessel elements in E.fra­ m; Wuhu, Anhui, HEFw 294, alt. 150 m. grans and E. japonica, or restricted to body Evergreen trees or shrubs mainly from of narrow vessel elements and tails of large South and Southwest China. vessel elementsin E. cavaleriei and E.dejlexa. Growth rings distinct to faint, marked by Gummy contents present in E. deflexa and rows of radially flattened latewood fibres, in E. japonica (HEFw 293 and CAFw 19781). few species also by differences in vessel fre­ Fibre-tracheids 910-1600 (640-2630) quency between latewood and subsequent um long, F/V ratio 1.3-2.1, medium thick­ earlywood. Vessels diffuse, 48-256/sq.mm to very thick-walled, with distinctly bordered (all above 140/sq.mm except in E. cavale­ pits of 4-7 um in diameter, in radial and tan­ riei), 73-98% (mostly above 90%) solitary, gential walls. Helical thickenings absent. remainder mainly in oblique, rarely tangential Parenchyma abundant, apotracheally dif­ and radial multiples of 2-3, round or oval to fuse, diffuse-in-aggregates and scanty para­ angular, tangential diameter 22-57 (13-90) tracheal, in 4-6 (2-1l)-celled strands. um, radial diameter 31-70 (15-135) um, Rays 8-12 (6-15)/mm, 1-3-seriate. Uni­ walls 1-3 um thick. Vessel element length seriate rays 2-5 (1-14) cells high, composed 480-930 (270-1290) um, L/D ratio 16-25. of square andupright cells; 2-3-seriate rays Perforations almost exclusively simple in 170-360 (50-820) um high, composed of oblique to almost horizontal end walls, but procumbent body cells and 1-3(-4) rows of sporadic scalariform and/or irregular multiple square to upright marginal cells (mainly het­ perforations with 2-5 bars also noted in E. erogeneous III, rarely heterogeneous 11). cavaleriei (CAFw 14), E. deflexa, E.jragrans Crystals sparse to abundant, prismatic, and E. japonica (Guangxi Forestry College sometimes elongated, large, present in eham­ s.n.). Intervessel pits nonvestured, alternate, bered axial parenchyma cells, one crystal in round to oval, 3-6 urn in diameter, with slit­ each enlarged, sclerified ehamber; chains of like apertures. Vessel-ray and vessel-paren­ up to 19 chambers. Pith flecks usually ab­ chyma pits similar to intervessel pits but half­ sent, but common in E.fragrans and one bordered and slightly smaller. Helical thick­ sample of E. japonica (FRIGw1060) . Trau­ enings well developed and closely spaced, matic gum ducts absent.

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Notes: 1. Moderately largevessels (200­ Malus Mill. (Figs. 7, 13,59; Tab1e 9) 300 um in diameter) in the genus reported by Material studied: M. baccata (L.) Borkh.: Metcalfe and Chalk (1950) are not present in Changbaishan, Jining, Qian Hong s.n., diam. these Chinese species. 13 mm; Changbaishan, Jining, CAFw 18070; 2. Moll and Janssonius (1914) reported Northeast China, Northeast Forestry Univer­ axial parenchyma very sparse in Eriobotrya sity s.n.; North China, CAFw 14225. - M. japonica. However, abundant diffuse and dif­ baccata (L.) Borkh. var. nigrecens: Zhong­ fuse-in-aggregates parenchyma, as in our tiaoshan, Shanxi, HEFw 121. - M. doumeri material, was recognised by Kanehira (1921a, (Bois) Chev.: Guanxi, Guanxi Forestry Col­ 1921b) . lege s.n. - M. hallianaKoehne : Hefei, Anhui,

Tab1e9. Variation in se1ectedwood anatomical features of Malus.

2 3 4 5 6 7 8 9 10 M. baccata Qian Hong s.n. 394 76 26 450 + 790 1.8 3-4 CAFw 18070 218 28 45 570 + 940 1.7 7-8 CAFw 14225 165 58 4;4 480 + 1080 2.2 6-7 var. nigrecens 235 61 39 510 1060 2.1 6-7 M. halliana 175 85 31 540 910 1.7 3-5 M. honanensis 168 74 35 420 + 850 2.0 3-5 + M. hupehensis Huang s.n. 385 51 25 420 + 780 1.9 4-5 HEFw 123 150 70 43 460 + 980 2.1 5-7 + M. manshurica 270 51 32 410 870 2.1 6-8 + M. melliana CAFw 15682 151 89 40 650 + 1080 1.7 7-8 + FRIGw 1814 208 60 34 650 1070 1.7 5-9 + HEFw 1377 183 80 32 610 + + 990 1.6 4-5 + M. pumila CAFw 13818 167 81 35 440 970 2.2 5-6 HEFw 1718 204 82 32 540 1080 2.0 4-6 HEFw 3570 155 88 32 430 + 930 2.2 5-6 CAFw 5652 158 73 39 450 980 2.2 5-6 M. rockii 73 84 42 500 1050 2.1 4-5 M. sibirica 293 73 29 440 + 890 2.0 6-7 M. spectabilis 268 70 30 500 + 1010 2.4 5-7 + M. toringoides 256 73 35 430 + 950 2.2 5-6 + M. yunnanensis 336 50 29 + 5-6

1: Vessel frequency (/sq.mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (um) ; 4: Average vessel element length (urn); 5: Multiple perforations present (+) or absent (-); 6: He1ical vessel wall thickenings present (+) or absent (-); 7: Average fibre-tracheid length (um); 8: F/V ratio; 9: Size of pits in fibre-tracheids (um); 10: Crystals present (+) or absent (-).

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S.Y. Zhang s.n., cult., diam. 18 mm, alt. 50 honanensis, M. hupehensis (Huang Chenglin m. - M. honanensis Rehd. : Zhongtiaoshan, s.n .), M. melliana (CAFw 15682 and HEFw Shanxi, HEFw 122. - M. hupehensis (pamp.) 1377) and M. yunnanensis, fine helical thick­ Rehd.: Qianliangfeng, Anhui, Huang Cheng­ enings often present in body of some narrow lin s.n., diam. 12 mm, alt. 300 m; Zhong­ vessel elements. Gummy contents present in tiaoshan, Shanxi, HEFw 123. - M. man­ some species. shurica (Maxim.) Kom.: Zhongtiaoshan, Fibre-tracheids 780-1080 (460-1390) Shanxi, HEFw 124. - M. melliana (Hand.« J.1m long, F/V ratio 1.6-2.4, medium thick­ Mazz.) Rehd.: Guangxi, CAFw 15682; Gu­ to very thick-walled, with distinctly border­ angdong, FRIGw 1814; Huangshan, Anhui, ed pits of 3-8 J.1ID in diameter, in both radial HEFw 1377, alt. 500 m. - M. pumila Mil!.: and tangential walls . Helical thickenings ab­ Xingjiang, CAFw 13818; Xingjiang, HEFw sent. 1718; Northeast China, HEFw 3570 ; Honan, Parenchyma moderately common to abun­ CAFw 5652. - M. rockii Rehd.: Tibet, CAFw dant, mainly apotracheally diffuse, but dif­ 19349. -M. sibirica Kom. : Northeast China, fuse-in-aggregates and scanty paratracheal HEFw 2397 . - M. spectabilis (Ait .) Borkh.: also present, in 3-5 (2-8)-celled strands. Hefei, Anhui, S.Y. Zhang s.n., cult., diam. Rays 8-12 (5-16)/mm, 1-3-seriate. 25 mm , alt. 50 m, - M. toringotdes (Rehd.) Unisenate rays 2-7 (1-22) cells high, com­ Hughes: Tibet, CAFw 19350. - M. yunnan­ posed of procumbent cells only or with one ensis (Franch.) Schneid.: Yunnan, CAFw row of square marginal cells; 2-3-seriate 67696. rays 160-320 (50-910) J.1m high, composed Deciduous, rarely semi-evergreen trees or of procumbent cells only, or rarely with one shrubs mainly from North China, widely cul­ row of square marginal cells (mainly homo­ tivated as fruit trees and ornamentals. geneous, but heterogeneous III also present Growth rings distinct, marked by rows of in most species, in two branch sampIes ofM. radially flattened latewood fibres, and also by hallianaand M. hupehensis, heterogeneous 11 differences in vessel frequency between late­ also present). wood and subsequent earlywood in M. hupe­ Crystals sparse to abundant in some spe­ hensis, M. pumi/a and M. rockii. Vessels dif­ cies (see Table 9), prismatic, large, present in fuse, 73-394/sq. mm (all above 150/sq. chambered axial parenchyma cells, one crys­ mm except in M. rockiii, 28-89% solitary tal in each enlarged, sclerified chamber; chains (all above 50% except in M. baccata, CAFw of up to 21 chambers. Pith flecks usually ab­ 18070), remainder mainly in oblique, rare­ sent, but rare to common in M. baccata var. ly tangential and radial multiples of 2-4, nigrecens, M. hupehensis (HEFw 123), M. oval, round to angular, tangential diameter manshurica, M. melliana (HEFw 1377), M. 25-45 (14-80) J.1m, radial diameter 32-70 rockii and M. yunnanensis. Traumatic gum (15-118) J.1ID, walls 1-2 J.1ID. Vessel element ducts absent. length 410-650 (210-1140) J.1m. LID ratio Notes: 1. Metealfe and Chalk (1950) stated 11-19. Perforations exclusively simple in ob­ that ring-porous or semi-ring-porous wood lique end walls, but in M. baccata, M. hupe­ OCCUfS in some species of the genus. But the hensis (HEFw 123), M. melliana (HEFw 13 Chinese species studied are all character­ 1377), M. pumila (HEFw 3570), M. sibirica, ised by diffuse-porous wood . Diffuse-porous M. spectabilis and M. toringoides, sporadic to weakly semi-ring-porous woods were, scalariform and/or irregular multiple perfora­ however, also reported by Schweingruber tions with 1-5 bars were also noted.lnterves­ (1990) for European species of Malus. seI pits nonvestured, alternate, mostly round, 2. Yang and Huang-Yang (1987) noted the occasionally polygonal, 5-10 J.1m in diam­ occurrence of libriform fibres in Malus for­ eter, with slit-like apertures. Vessel-ray and mosana. No libriform fibres, however, were vessel-parenchyma pits usually with reduced found in OUf material. borders and smaller, 3-5 J.1m in diameter. 3. Multiple perforations in the genus were Helical thickenings mostly absent, but in M. also recorded by Brazier and Franklin (1961).

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Table 10. Variation in selected wood anatomical features of Photinia.

2 3 4 5 6 7 8 9 10 11 12 13 Evergreen group (A)

P. davidsoniae Zhang s.n, 186 95 29 560 + + 960 1.7 15 2 0.3 + FRIGw 1179 155 86 31 660 + 1150 1.7 12 2 0.2 + HEFw 2674 180 82 37 660 + 1150 1.8 + 12 3 0.2 + P. glabra HEFw 596 200 97 24 480 + 770 1.6 10 2 0.2 + Guizhou s. n. 167 80 36 620 + + 1180 1.9 + 11 2 0.2 - HEFw 267 257 92 25 510 + + 990 1.9 + 14 2 0.2 + CAFw 13053 112 93 31 640 + 1250 2.0 + 13 2 0.3 + P. prunifolia 161 93 30 620 + 930 1.5 + 12 2 0.2 + P. serrulata HEFw 2000 259 90 24 470 + + 810 1.7 + 12 2(-3) 0.2 + FRIGw 1843 165 94 29 490 + 900 1.9 + 13 2(-3) 0.2 +

Deciduous group (B)

P. beauverdiana 120 90 44 600 + + 1190 2.0 + 6 4(-5) 0.3 + P. bentham iana 104 72 47 750 + - 1310 1.7 7 4(-5) 0.4 + P. schneideriana 100 94 45 670 + 1560 2.3 + 7 4(-5) 0.4 + P. villosa 145 85 40 650 + + 1210 1.8 + 9 4(- 5) 0.5 +

1: Vessel frequency (/ sq. mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (11m); 4: Average vessel element length (um); 5: Multiple perforations present (+) or absent (-); 6: Helical vessel wall thickenings present (+) or absent (-); 7: Average fibre-tracheid length (um); 8: F/V ratio; 9: Helical thickenings present in fibre-tracheids (+) or absent (-); 10: Ray frequency (jmm); 11: Width of the widest rays (number of cells); 12: Average multi seriate ray height (11m); 13: Crystals present (+) or absent (-).

Photinia Lind!. (Figs. 15, 17,43,60; Table Anhui, HEFw 596 ; Guangzhou, Guizhou 10) Forest Institute s.n.; Anqing, Anhui, HEFw This genus, mainly from South, South­ 267, alt. 300 m; Fujian, CAFw 13053; Gu­ west and East China , consists of both ever­ angxi , Guangxi Forestry College s.n. - P. green and deciduous trees or shrubs. Wood prunifolia (Hook. & Am .) Lindl.: Guang­ anatomical differences between the evergreen dong , FRIGw 1257. - P. serrulata Lindl. : and deciduous species are appreciable. Thus Huang shan, Anhui, HEFw 2000, alt. 500 m; two separate descriptions are given. Guangdong, FRIGw 1843; Wuhu, Anhui, HEFw 295, alt. 200 m; Hefei, Anhu i, S.Y. Evergreen group (Group A) Zhang s.n., cult., alt. 50 m. Material studied: P. davidson iae Rehd. & Growth rings distinct to faint, marked by Wils.: Hefei, Anhui , S.Y. Zhang s.n ., cult. , rows of radially flattened latewood fibres. alt. 50 m; Guangdong, FRIGw 1179; Hunan , Vessels diffuse , (l 12-) 155-259/sq.mm, HEFw 2674. - P. glabra (Thunb.) Maxim .: 80 -97% (mostly above 90 %) solitary, re-

Downloaded from Brill.com10/09/2021 05:06:33PM via free access 58 IAWA Bulletin n.s., Vol. 13 (1),1992 mainder mainly in oblique, rarely tangential P. prunifolia, small to large. Traumatic gum and radial multiples of 2(-4), round, oval to ducts absent. angular, tangential diameter 24-37 (12-60) 11m, radial diameter 29-51 (15-75) 11m, Deciduous group (Group B) walls I-211m thick. Vessel element length Material studied: P. beauverdiana Schneid.: 470-660 (270- 960) 11m. LID ratio 17-21. Yaan, Sichuan, Sichuan Forestry Institute Perforations almost exclusively simple in s.n.; Southwest China, HEFw 1144. - P. oblique end walls, but sporadic scalariform benthamiana Hance: Guangdong, FRIGw and/or irregular multiple perforations with 1939. -Po schneideriana Rehd. & Wils.: Gu­ 2 - 5 bars occur in all species except in P. angdong, FRIGw 1206. -Povillosa (Thunb.) prunifolia where perforations are exclusively DC.: Sichuan, CAFw 7009. simple. Intervessel pits nonvestured, alter­ Growth rings distinct to faint, marked by nate, round to oval, 3-7 um in diameter, usu­ rows of radially flattened latewood fibres. ally with slit-like apertures . Vessel-ray and Vessels diffuse, 100-145/sq.mm, 72-94% vessel-parenchyma similar to intervessel pits solitary, remainder mainly in oblique, rarely but half-bordered and slightly smaller. Heli­ tangential and radial multiples of 2-4, round, cal thickenings mostly weil developed and oval to angular, tangential diameter 40-47 closely spaced, occasionally widely spaced in (20-70) 11m, radial diameter 48-62 (22-90) R. davidsoniae (HEFw 2674), throughout 11m, walls I-311m thick. Vessel element body of vessel elements, but in a few species, length 600-750 (270-1390) 11m. L/D ratio fine helical thickenings only present in body 14-16. Perforations almost exclusively sim­ of narrow vessel elements. Gummy contents ple in oblique to almost horizontal end walls, present in some vessels of P. davidsonia but scalariform and irregular multiple perfo­ (FRIGw 1179) and P. prunifolia. rations recorded in all species except in P. Fibre-tracheids 770-1250 (330-1580) 11m schneideriana. Intervessel pits nonvestured, long, F/V ratio 1.5-2.0, medium thick- to alternate, round, 3-7 11m in diameter, with very thick-walled, with distinctly bordered slit-like apertures. Vessel -rays and vessel­ pits of 3-6lJ.ffi in diameter, in radial and tan­ parenchyma pits similar to intervessel pits gential walls. Helical thickenings fine, pres­ but half-bordered and slightly smaller. Heli­ ent in most species, but absent in P. david­ cal thickenings weil developed and closely soniae (Zhang, S.Y. s.n. and FRIGw 1179) spaced, throughout body of vessel elements , and P. glabra (HEFw 596). but the thickenings not observed in P. ben­ Parenchyma abundant, apotracheally dif­ thamiana. Gummy contents absent. fuse, diffuse-in-aggregates and scanty para­ Fibre-tracheids 1190-1560 (880- 2060) tracheal, in 2-5 (2-8)-celled strands. 11m long, F/V ratio 1.7-2.3, medium thick­ Rays 10-15 (5-16)/mm, 1-2(-3)-seri­ to very thick-walled, with distinctly bordered ate. Uniseriate rays 2-3 (1-9) cells high, pits of 4-7 um in diameter, in radial and tan­ composed of square (occasionally to weakly gential walls. Helical thickenings fine, present procumbent) cells and upright cells; multi­ in all species except in P. benthamiana. seriate rays 150-300 (30-600) 11m high, Parenchyma abundant, apotracheally dif­ composed of procumbent body cells and 1-3 fuse, diffuse-in-aggregates and scanty para­ rows of square to upright marginal cells (het­ tracheal, in 3-5 (2-1O)-eelled strands. erogeneous III & II). Rays 6-9 (5-1O)/mm, 1-4(-5)-seriate, Crystals mostly abundant, but infrequent of rwo sizes. Uniseriate rays 3-4 (1-10) in P. glabra (CAFw 13053), prismatic, occa­ cells high, composed of square (to weakly sionally elongated in a few species, typically procumbent) cells and upright cells; multi­ large, present in chambered axial parenchyma seriate rays 280-490 (60-1200) 11m high, cells, one crystal in each enlarged, usually composed of procumbent body cells and sclerified chamber; chains of up to 23 cham­ 1-2(-3) rows ofmarginal cells (predominant­ bers, crystals not observed in P. glabra ly heterogeneous III, rarely heterogeneous II, (Guizhou Forestry Institute s.n.). Pith flecks but homogeneous rays occasionally also common in P. davidsoniae (HEFw 2674) and present).

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Crystals sparse to abundant in all species, Pyracantha Roem. prismatic, large, occasionally elongated, in Material studied: Pifortuneana (Maxirn.) chambered axial parenchyma cells, one crys­ Li: Hefei, Anhui, S.Y. Zhang s.n., cult., tal in each enlarged chamber; chain of 14 diam. 20 mm, alt. 50 m. chambers. Pith flecks common in P. schnei­ Evergreen shrubs or small trees mainly deriana and P. villosa, large, in tangential from Southwest, Northwest and South China. bands. Traumatic gum ducts absent Growth rings distinct, marked by 2-6 Notes : 1. The differences between the rows of radially flattened latewood fibres. evergreen and deciduous species are restricted Vessels diffuse, 224/sq.mm, 88% solitary, to quantitative features. The deciduous group, remainder in oblique, radial and tangential as shown in Table 10, has Ionger and wider multiples of 2-3, round, oval to angular, vessel elements, a lower vessel frequency , tangential diameter 32 (17-50) 11m, radial and significantly longer fibre-tracheids. In diameter 40 (20 -55) 11m, walls I-211m addition, the deciduous species have a lower thick. Vessel element length 290 (210-390) ray frequency, and rays are wider and higher. 11m. LID ratio 9. Perforations almost exclu­ Similar results were reported by Kanehira sively simple in oblique end walls, but spor­ (1921b): the differences in quantitative fea­ adic scalariform perforations with up to 5 tures between deciduous P. variabilis and bars also noted. Intervessel pits nonvestured , evergreen P. glabra are generally in accor­ alternate, round to oval, 5-7 um in diameter, dance with the ones recognised between the with slit-like apertures. Vessel-ray and ves­ two groups in the present study. In his study sel-parenchyma pits similar to intervessel on Formosan woods (Kanehira 1921a), sim­ pits but half-bordered and slightly smaller. ilar differences between deciduous P. serru­ Helical thickenings fine and closely spaced, lata and evergreen P. taiwanensis were also present in most vessel elements. Gummy reported. Photinia taiwanensis is character­ contents abundant, present in most vessels. ised by 1-8-seriate nearly homogeneous rays, Fibre-tracheids 590 (390-740) 11m long, significantly lower vessel frequency, wider F/V ratio 2.0, medium thick-walled, with dis­ vessel elements and longer fibres than P. ser­ tinctly bordered pits of 3-5 11m in diameter, rulata which has 1-2-seriate heterogeneous in radial and tangential walls. Helical thicken­ rays. Lu et al. (1991) also found significant ings present in almost all fibre-tracheids. differences in leaf epidermal characters be­ Parenchyma abundant, apotracheally dif­ tween the deciduous and evergreen species fuse and scanty paratracheal , in 4 (3-8)-cell­ groups. ed strands. 2. Metcalfe and Chalk (1950) mentioned Rays 13 (1O-16)/mm, 1-3-seriate . Uni­ that some species in Photinia are ring-porous seriate rays 6 (1-8) cells high, composed of or semi-ring-porous. However, all species square to weakly procumbent and upright studied here are diffuse-porous. Diffuse­ cells ; 2-3-seriate rays 280 (70-610) 11m porous woods in Photinia species were also high, composed of procumbent body cells reported by Cheng (1980). and 1-4(-6) rows of square to upright mar­ 3. Occurrence of very fine septate fibres ginal cells (heterogeneous III and 11). in Photinia lindleyana was documented by Crystals are sparse, prismatic, sometimes Metcalfe and Chalk (1950), but we did not elongated, medium-sized to large, in cham­ observe septate fibres in the Chinese species. bered axial parenchyma cells, one crystal in 4. Vessel frequencies of 30-40/sq.mm each enlarged and sclerified chamber; chains reported for the genus by Metcalfe and Chalk ofup to 6 chambers. Pith flecks and traumat­ (1950) are much lower than our values. ic gum ducts absent. 5. The deciduous species of Photinia are Notes : 1. The drawings by Greguss (1959) the only ones in the Maloideae with rays of indicate the absence of helical thickenings in two sizes and wider than 3(-4)-seriate. fibres, but they are present in almost all fibre­ 6. Moll and Janssonius (1914) reported tracheids in the Chinese species. Dur descrip­ axial parenchyma as very sparse in Photinia. tion closely resembles that of P. crenulata However, this is not the case in our material. from the Himalayas by Suzuki et al. (1991).

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Table 11. Variation in selected wood anatomical features of Pyrus.

1 2 3 4 5 6 7 8 9 10 11 12 P. betulaefolia 340 67 27 480 5-7 920 1.9 3 HelHo - P. calleryana HEFw 3710 170 65 35 530 6-10 940 1.8 + 3 Ho HEFw 1661 298 53 29 590 + 6-7 940 1.6 + 3 Ho P. pashia 200 90 31 540 5-9 1110 2.1 + 2 Ho + P. pyrifolia Zhang s.n . 278 55 31 460 6-7 780 1.7 3 HelHo - 367 128 71 47 360 6-8 860 2.4 + 3 He/Ho - var. (culta) 167 81 37 430 5-7 910 2.1 + 2(-3) Ho + P. serrulata HEFw 329 210 59 38 590 + 6-7 1060 1.8 + 3 Ho + FRIGw 1793 142 75 41 + 5-10 + 2 HelHo - HEFw 785 169 79 33 530 5-8 950 1.8 - 2(-3) Ho + P. ussuriensis CAFw 5928 333 34 26 560 6-8 940 1.7 + 3(-4) He/Ho + Northeast s.n . 195 76 34 6-10 - 2(-3) Ho CAFw 18077 288 68 34 ;60 + 9-11 1200 2.1 + 2 Ho P. xerophila 188 85 35 500 7-10 980 2.0 - 2(-3) Ho 1: Vessel frequency (/sq.mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (IJ.m); 4: Average vessel element length (um); 5: Multiple perforations present (+) or absent (-); 6: Size of intervessel pits (um); 7: Average fibre-tracheid length (um); 8: F/V ratio; 9: Helical thickenings present in fibre-tracheids (+) or absent (-); 10: Width of the widest rays (number of cells); 11: Rays homocellular only (Ho), or heterocellular III also present (He/Ho) ; 12: Crystals present (+) or absent (-) .

2. Homogeneous rays were reported by uan, HEFw 785. - P. ussuriensis Maxim.: Greguss (1959) in Pyracantha coccinea, Sichuan, CAFw 5928; Northeast China, which is different from the situation in P. Northeast Forestry University s.n .; Wuhu, fortuneana . Anhui, HEFw 418, alt. 100 m; Jining, CAFw 18077. - P. xerophila Yu: Zhongtiaoshan, Pyrus L. (Table 11) Shanxi, HEFw 136. Material studied: P. betulaefolia Bunge: Deciduous trees or shrubs, rarely semi­ Anhui, HEFw 308. - P. calleryana Decne.: evergreen trees mainly from North China, Huoshan, Anhui, HEFw 3710, alt. 150 m; widely cultivated as fruit trees and omamen­ Jixi, Anhui, HEFw 3185, alt. 550 m; Jiang­ tals. xi, HEFw 1661; Guangdong, FRIGw 1369. Growth rings distinct, marked by 1-4 - P. pashia Buch-Harn. ex D. Don.: Gui­ rows of radially flattened latewood fibres, zhou, Guizhou Forestry Institute s. n. - P. and also by differences in vessel frequency pyrifolia (Burm. f.) Nakai: Hefei, Anhui, between latewood and subsequent earlywood S.Y. Zhang s.n., cult., alt. 50m; Kumming, in P. calleryana (HEFw 1661) and P. ussuri­ Yunnan, Yunnan Academy of Forestry 367, ensis. Wood usually diffuse-porous (but in alt. 1900 m. - P. pyrifolia (Burm. f.) Nakai P. ussuriensisranging to weakly semi-ring­ var.: Guangdong, FRIGw 1604. - P. serru­ porous). Vessels 128-340/sq.mm, 34-90 lata Rehd.: Huoshan, Anhui, HEFw 329, % (mostly above 50%) solitary, remainder alt. 250 m; Guangdong, FRIGw 1793; Sich- mainly in oblique, rarely tangential and radial

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 61 multiples of 2-6, round, oval to angular, occasionally with Ollerow of square marginal tangential diameter 26-47 (20-70) lUD, ra­ cells (predominantly homogeneous, but het­ dial diameter 33-70 (20-103) lJ.Il1, walls 1-3 erogeneous m occasionally present in some 11m thick. Vessel element length 360-590 species). (210-830) 11m. LID ratio (8-)15-22. Per­ Crystals sparseto common in P. pashia,P. forations mostlyexclusively simple in oblique pyrifolia var., P. serrulaia (HEFw 329 and end walls, but sporadic scalariform and/or HEFw 785) and P. ussuriensis (CAFw 5928), irregular multiple perforations with 1-4 bars prismatic, occasionally elongated in P. ussur­ occur in P. cal/eryana (HEFw 1661), P. ser­ iensis, large, in chambered axial parenchyma rulata (HEFw 329 and FRlGw 1793) and P. cells, one crystal in each enlarged, sclerified ussuriensis (CAFw 18077) . Intervessel pits chamber; chains of up to 12 chambers, crys­ nonvestured, alternate, round to oval , 5-11 tals absent in other species or specimens. Pith lUD in diameter, with slit-like apertures, aper­ flecks usually absent, but occasionally tures occasionally coalescent in some species. present in P. pyrifolia (Yunnan Academy of Vessel-ray and vessel-parenchyma pits sim­ Forestry 367). Traumatic gum ducts absent ilar to intervessel pits but half-bordered and Note: Absence of helical thickenings in smaller, 4-6 lUD in diameter. Helical thick­ vessel elements ofPyrus pashia wasreported enings well developed and closely to mod­ by Luo (1989). However, fine thickenings erately widely spaced, throughout body of were noted in narrow vessel elements and tails vessel elements in P. cal/eryana (HEFw 3710 of large vessel elements in the species we and HEFw 1661), P. pyrifolia, P. serrulata examined. (FRlGw 1793 and HEFw 329) and P. ussur­ iensis (CAFw 5928); or fine spiral thicken­ Raphiolepis Lindl. (Fig. 27; Table 12) ings restricted to body of narrow vessel Material studied: R. ferruginea Metealf: elements and tails of large vessel elements in Fujian, HEFw 3861 ; Guangdong, FRIGw P. betulaefolia, P. pashia, P. serrulata (HEFw 1302. - R. indica (L.) Lindl.: Qianliangfeng, 785), P. ussuriensis (Northeast Forestry Uni­ Anhui, Huang Chenglin s. n., diam. 8 mm, versity s.n.) and P. xerophila. Gummy con­ alt. 300; Hainan, South China Agricultural tents absent. University 831; Guangdong, Guizhou Fores­ Fibre-tracheids 780-1200 (520-1650) try Institute s.n .; Guangdong, FRlGw 1150. 11m long, F/V ratio 1.6-2.4, very thin- to -R.lanceolata Hu: Guangxi, CAFw 16126. very thick-walled, with distinctly bordered -R. salicifolia Lindl.: Guangdong, FRlGw pits of 4-8 11m in diameter, numerous in 1779 . radial walls and usually in tangential walls as Evergreen shrubs or small trees mainly well (except in P. cal/eryana, HEFw 3710) . from South China. Helical thickenings fine to conspicuous, Growth rings distinct to faint, marked by present in most species, but absent in some rows of weakly radially flattened latewood specimens, e.g., P. betulaefolia,P. pyrlfolia fibres, and in a few samples also by differ­ (Zhang s.n.), P. serrulata (HEFw 785), P. ences in vessel frequency between latewood ussuriensis (Northeast Forestry University and subsequent earlywood. Vessels diffuse, s.n.) and P. xerophila. 118-238(-319)/sq.mm, 71-98% solitary, Parenchyma fairly common to abundant, remainder mainly in oblique, rarely tangential mainly apotracheally diffuse, but diffuse­ and radial multiples of 2-3, round or oval to in-aggregates and scanty paratracheal also angular, tangential diameter 16-40 (12-57) present in most species, in 3-6 (2-9)-celled lUD, radial diameter 17-45 (13-70) 11In, walls strands. 1-2 um thick. Vessel element length 370­ Rays 7-13 (5-16)/mm, 1-3(-4)-seriate. 860 (210-1250) lUD. LID ratio (13-)19-27. Uniseriate rays 2-6 (1-16) cells high, com­ Perforations exclusively simple in oblique posed of procumbent cells only, or occasion­ end walls, but in R. indica (FRIGw 1150 , ally with one row of square marginal cells; South China Agricultural University 831) and 2-3-serlate rays 180-280 (50-530) 11m R. salicifolia, sporadic scalariform and/or high, composed ofprocumbent cells only, or irregular multiple perforations with 2-4 bars

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Tab1e 12. Variation in se1ected wood anatomical features of Raphiolepis.

2 3 4 5 6 7 8 R. ferruginea HEFw 3861 118 32 860 1420 1.6 0.4 FRIGw 1302 238 34 770 1090 1.4 0.3 R. indica Huang s.n, 319 16 510 760 1.5 0.3 831 143 40 760 + 1080 1.4 0.4 + Guizhou s.n, 197 29 370 800 2.1 0.2 FRIGw 1150 176 34 670 + 1140 1.7 0.3 + Ri lanceolata 200 30 640 1030 1.6 0.2 + R. salicifolia 187 34 790 + 1120 1.4 0.3 +

1: Vesse1 frequency (/sq.mm); 2: Average tangential vessel diameter (um); 3: Average vessel element length (um); 4: Multiple perforations present (+) or absent (-); 5: Average fibre-tracheid length (um) ; 6: F/V ratio; 7: Average multiseriate ray height (mm); 8: Crystals present (+) or absent (-). also noted. Intervessel pits nonvestured, alter­ Crystals sparse to abundant, prismatic, nate, round to oval, 3-6 11m in diameter, large, in chambered axial parenchyma cells, with slit-like apertures. Vessel-ray and ves­ one crystal in each enlarged sclerified cham­ se1-parenchyma pits simi1ar to intervesse1 ber; chains of up to 17 chambers, crystals not pits but half-bordered and slightly smaller. observed in Riferruginea and R. indica (Gui­ Helical thickenings weil developed and close­ zhou Forest Institute s.n. and Huang s.n .). ly spaced, throughout body of most vessel Pith flecks common in R. indica (except in elements. Gummy contents present in a few one sampie) andR.lanceolata. Traumatic gum vessels of two specimens ofR. indica (South ducts absent. China Agricultural University 831 and Huang Note: Rays in Raphiolepis, as weil as in Chenglin s.n.) and R. salicifolia. Stranvaesia, are the most heterogeneous (in­ Fibre-tracheids 760-1420 (510-1600) um cluding Kribs' type I) in the subfamily. Het­ long, F/V ratio 1.4-2.1, medium thick- to erogeneous I type is sometimes present in the very thick-wal1ed, with distinctly bordered two genera . pits of (4-)5-6 um in diameter, in radial and tangential walls . Helical thickenings usually Sorbus L. (Figs . 14, 16, 18, 23, 29, 39, 48; fine, present in a few to most fibre-tracheids Table 13) of all species . Materials studied: S. alnifolia (Sieb. & Parenchyma abundant, apotracheally dif­ Zucc .) K. Koch: Zhongtiaoshan, Shanxi, fuse, diffuse-in-aggregates and scanty para­ HEFw 138; Shandong, HEFw 1360; North­ tracheal, in 3-5 (2-11)-eelled strands. east China, CAFw 13729; Guangxi, CAFw Rays 10-13 (7-19)/mm, 1-2-seriate. 5588; Dongling, Hobai, CAFw 5315; Honan, Uniseriate rays 2-4 (1-14) cells high, com­ CAFw 5499. - S. alnifolia (Sieb.& Zucc.) posed of upright cells only, or occasionally var. lobulata Rehd .: Zhongtiaoshan, Shanxi, with square cells; biseriate rays 170-410 HEFw 140. - S. amabilisCheng ex Yu: Qian­ (40-830) 11m high, composed of procum­ liangfeng, Anhu i, Huang Chanlin s. n., diam. bent body cells and 1-6 rows ofsquare to up­ 12 mm, alt. 350 m. - S. aria (L.) Crantz: right marginal cells [heterogeneous III & II; Hubei, CAFw 9798. -S. caloneura (Stapf) type I also present in Rcferruginea,R. indica Rehd.: Guizhou, Guizhou Forestry Institute (South China Agric. Univ. 831 and Huang s.n. - S. commixta Hedl.: locality & col!. Chenglin s.n .) and R. salicifolia]. unknown.-S. coronata (Card .) Yu & Tsai :

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Table 13. Variation in selected wood anatomical features of Sorbus. 2 3 4 5 6 7 8 9 10 11 12 S. alnifolia HEFw 138 175 66 33 590 + 5-8 1110 1.9 + 3 + + HEFw 1360 242 71 37 600 + 4-5 1040 1.7 2 CAFw 13729 209 54 44 770 + 4-7 1170 1.5 + 2 CAFw 5588 324 54 31 670 + 5-7 940 1.4 2 + CAFw 5315 320 58 36 600 + 5-7 1040 1.7 2 var. lobulata 227 67 36 340 - 5-6 870 2.5 2 S. amabilis 268 66 24 380 - 5-6 730 1.9 2 + + S. aria 138 89 33 590 + 4-6 1150 1.9 3 + S. caloneura 208 67 33 640 + 5-6 1120 1.8 + 3 + S. coronata HEFw 1187 161 64 46 610 + 6-8 1120 1.8 3 CAFw7601 171 80 34 550 + 7-9 1090 2.0 2 + S. discolor HEFw 139 216 74 32 470 - 8-10 860 1.8 + 2(-3) - CAFw 5400 233 57 29 420 + 5-7 800 1.9 + 2(-3) + + S. dunnii 109 79 45 760 + 5-7 1040 1.8 + 2(-3) + S. folgneri HEFw 2657 139 84 40 760 - 5-6 1310 1.7 + 3 + 127 245 59 35 770 - 4-6 1240 1.6 3 CAFw 10963 113 73 47 730 - 5-8 1440 2.0 3 Guangxi s.n, 142 74 43 810 + 4-5 1240 1.5 3 + S. hemsleyi HEFw 874 106 98 39 610 + 3-5 1020 1.7 + 3 + 393 209 68 35 600 - 6-8 1140 1.9 2 + FRIGw 1249 140 78 35 700 - 5-7 1220 1.7 + 3 + S. koehneana HEFw 141 263 62 30 420 + 5-6 790 1.9 + 2 + + CAFw 509 232 77 30 430 + 5-7 780 1.8 + 2 + S. microcarpa 183 89 32 590 + 4-5 1150 2.0 4 + S. oligodonta 477 64 26 640 + 7-10 1020 1.6 2 + S. pohuashanensis 215 63 32 440 + 4-6 820 1.9 3 + S. prattii 860 208 90 28 430 + 5-7 720 1.7 2 CAFw7622 199 63 36 610 + 6-8 1040 1.7 2 CAFw 19321 330 68 24 590 + 5-8 1020 1.7 - 2(-3)- + var. tatsienensis 189 67 39 350 5-8 1150 3.3 4 S. pteridophylla 175 70 38 550 + 7-10 950 1.7 + 2 + S. reducta 218 74 30 420 + 5-7 850 2.0 2 + S. sargentiana 150 72 45 670 + 5-8 1080 1.6 2 + S. tianschanica 327 63 31 510 + 4-6 900 1.8 2 S. vilmorinii 127 80 32 580 + 5-10 990 1.7 2 S. wallichii 219 65 33 610 - 8-10 1120 1.8 2 S. wilsoniana 207 63 33 600 + 5-6 1060 1.8 3 + 1: Vessel frequency Usq.mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (um); 4: Average vessel element length (um); 5: Multiple perforations present (+) or absent (-); 6: Size of intervessel pits (um); 7: Average fibre-tracheid length (um); 8: F/V ratio; 9: Helical thickenings present in fibre-tracheids (+) or absent (-); 10: Width of the widest rays (number of cells); 11: Crystals present (+) or absent (-); 12: Wood diffuse (-) or tending to be semi-ring-porous (+).

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Southwest China, HEFw 1187; Taiwan, donta), the frequency often decreases from CAFw 7601; Shaping, Sichuan, CAFw 379. earlywood to latewood in some species, 54­ - S. discolor (Maxim.) Maxim.: Zhongtiao­ 98% solitary (mostly 63-90%), remainder shan, Shanxi, HEFw 139; Wutashan, Shanxi, mainly in oblique, rarely tangential and radial CAFw 5400. - S. dunnii Rehd.: Guizhou, multiples of 2-4, oval, round to angular, Guizhou Forest Institute s.n. - S. folgneri tangential diameter 24-47 (13-68) um, ra­ (Schneid.) Rehd.: Hunan, HEFw 2657; Yi­ dial diameter 35-62 (15-105) um, walls nian, Yunnan, Yunnan Academy of Forestry 1-3 um thick. Vessel element length 340­ 127, alt. 2000 m; Sichuan , CAFw 10963; 810 (190-1360) um. LID ratio (10-)13-24. Guangxi, Guangxi Forestry College s.n. ­ Perforations almost exclusively simple in S. hemsleyi (Schneid.) Rehd.: Sichuan, oblique end walls, but sporadic scalariform HEFw 874; Lijiang, Yunnan, Yunnan Acade­ and/or irregular multiple perforations with my of Forestry 393, alt. 3000 m; Guangdong, 2-5 bars occur in most species, only in S. FRIGw 1249. - S. koehneana Schneid.: alnifolia var. lobulata, S. amabilis, S. dis­ Zhongtiaoshan, Shanxi, HEFw 141; Taibai­ color (HEFw 139), Sifolgneri (HEFw 2657, shan, Shanxi, CAFw 509. - S. microcarpa Yunnan Academy ofForestry 127 and CAFw Pursh: Sharping, Sichuan, CAFw 346.­ 10963), S. hemsleyi (Yunnan Academy of S.oligodonta (Card.) Hand.-Mazz.: Tibet, Forestry 393 and FRIGw 1249), S. prattii CAFw 19322. - S. pohuashanensis(Hance) var. tatsienensis and S. wallichii, perforations Hedl.: Dongling, Hobai, CAFw 5317. - S. are exclusively simple. Intervessel pits non­ prattii Koehne: Zhongdian, Yunnan, Yunnan vesmred, alternate, round to oval, 4-8(-10) Academy of Forestry 860, alt. 3500 m; Tai­ um in diameter, with slit-like apertures, aper­ wan, CAFw 7622; Tibet, CAFw 19321. - S. tures occasionally coalescent in a few spe­ prattii Koehne var. tatsienensis (Koehne) eies. Vessel-ray and vessel-parenchyma pits Schneid.: Tibet, CAFw 10660 - S. pterido­ similar to intervessel pits but half-bordered phylla Hand.-Mazz.: Lijiang, Yunnan, Yun­ and slightly smaller. Helical thickenings weIl nan Academy of Forestry 404, alt. 3000 m. ­ developed and closely spaced, throughout S. reducta Diels: Zhongdian, Yunnan, Yun­ body of vessel elements. Gummy contents nan Academy ofForestry 861, alt. 3500 In.­ present in a few species. S. sargeniiana Koehne: Shaping, CAFw 291. Fibre-tracheids 720-1440 (440-1920) - S. scalaris Koehne: Shaping, Sichuan, um long, F/V ratio 1.4-2.5(-3.3), medium CAFw 310. - S. tianschanica Rupr.: Xinji­ thick- to very thick-walled, with distinctly ang, HEFw 1501. - S. vilmorinii Schneid.: bordered pits of 4-7 um in diameter, in radial location unknown, CAFw 19342. - S. wal­ and tangential walls. Helical thickenings fine lichii (Hook.f.) Yu: Jianchuan, Yunnan, Yun­ to conspicuous, present in S. alnlfolia (HEFw nan Academy of Forestry 636, alt. 2000 m, 138 and CAFw 13729), S. caloneura, S. dis­ - S. wilsoniana Schneid. : Sichuan, CAFw color, S. dunnii, S. folgneri (HEFw 2657), 6292. S. hemsleyi (HEFw 874 and FRIGw 1249), Deeiduous trees or shrubs mainly from S. koehneana and S. pteridophylla, but heli­ Southwest China. cal thickenings not observed in the remaining Growth rings distinct, marked by rows of species or sampies. radially flattened latewood fibres, and also by Parenchyma moderately common to abun­ differences in vessel frequency between late­ dant, mainly apotracheally diffuse, but dif­ wood and subsequent earlywood in some fuse-in-aggregates and scanty paratracheal species. Wood diffuse-porous, rarely ranging also present in most species, in 3-6 (2-11)­ to semi-ring-porous in S. alnifolia (HEFw celled strands. 138), S. amabilis, S. caloneura, S. discolor Rays 6-16 (3-20)/mm, 1-3(-4)-seriate (CAFw 54(0), S.folgneri (Guangxi Forestry (mostly 1-2(-3)-seriate). Uniseriate rays 3-8 College s.n.), S. koehneana, S. pohuasha­ cells high, composed of procumbent cells on­ nensis, S. prattii (CAFw 19321), S. reducta ly, or with one row of square marginal ceIls; and S. wilsoniana. Vessels 106-477/sq.mm 2-3(-4)-seriate rays 130-350 (50-820) (all below 330/sq.mm except in S. oligo- um high, composed of procumbent cells only,

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 65 or with 1(-2) row(s) of square marginal cells walled, with distinctly bordered pits of 4-6 (mostly homogeneous , rarely heterogeneous um in diameter, in radial and tangential walls. III). Helical thickenings fine, present in a few Crystals sparse to abundant in some spe­ fibre-tracheids. cies (see Table 13), prismatic, sometimes Parenchyma moderately abundant, pre­ elongated in a few species, large, present in dominantly apotraeheally diffuse, morerarely chambered axial parenchyma cells,one crys­ diffuse-in-aggregates and scanty paratracheal, tal in each enlarged, sclerifiedchamber;chains in 3 (2-7)-celled strands. of up to 20 chambers. Pith flecks rare to com­ Rays 14 (1l-18)/mm, 1-2-seriate. Uni­ mon in some species. Traumatic gum ducts seriate rays 4 (1-11) cells high, composed of absent. upright cells only, or rarely with square cells; Notes: 1. This study of over 20 species biseriate rays 250 (70-420) um high, com­ of Sorbus, covering all three sections Aria, posed ofprocumbent body cells and 1-4(-6) Microme/es and Sorbus recognised by Yu rows of square to upright marginal cells (1974), indicates that the genus is wood ana­ (mainly heterogeneous mand Il, rarely het­ tomically highly uniform. Sorbus commixta, erogeneous I). Microme/es alnifolia and M. japonica were Crystals common, prismatic, occasionally studied by Kanehira (1921a) and no appreci­ elongated, medium-sized to large, present in able differences were recognised between the chambered axial parenchyma cells, one crys­ Microme/es species (now Sorbus) and Sorbus tal in each enlarged, weakly sclerified cham­ commixta. ber; chains of up to 23 chambers. Prismatic 2. Turnanin (1949) also recognised the or fragmented crystals also occasionally pres­ variation from diffuse-porous to semi-ring­ ent in non-enlarged chambers, one to several porous in nine Sorbus species he studied. Met­ crystals per chamber. Pith flecks common, in calfe and Chalk (1950) recorded that some tangentialbands. Traumatic gum ducts absent, species in Sorbus are ring-porous or semi­ Notes: 1. This is the first wood anatomical ring-porous. descrlption of Stranvaesia. 2. Kalkman (1973) reduced the genus to Stranvaesia Lindl. Photinia. Wood anatomically there are no Material studied: S. davidiana Decne.: Si­ significant differences between Stranvaesia chuan, CAFw 357, diam. 30 mm. and Photinia s.str., so that their merger is not Evergreen trees or shrubs mainly from in conflict with the anatomical evidence. Leaf Southwest China, often cultivated as orna­ anatomy also supports this (Lu et a/. 1991). mentals. Growth rings distinct, marked by rows of Subfamily m: Rosoideae Focke radially flattened latewood fibres. Vessels dif­ fuse, 419/sq.mm, 77% solitary, remainder Kerria Oe. mainly in oblique and tangential multiples of Material studied: K. japonica (L.) DC.: 2-3, mostly angular, tangential diameter 22 Huangshan, Anhui, Liu Xiumei s.n., diam. (16-36) um, radial diameter 26 (17-45) 10 mm, alt. 600 m; Qingliengfeng, Anhui, um, walls 1-2 um thick. Vessel element Huang Chenglin s.n., diam. 12 mm, alt. length 530 (230-760) um, LID ratio 24. Per­ 550m. forations exclusively simple in oblique end Monotypic genus, deciduous shrubs from walls. Intervessel pits nonvestured, alternate, Northwest to southeastern China, widely cul­ round, 3-5 um in diameter, with slit-like tivated as ornamentals. apertures. Vessel-ray and vessel-parenchy­ Growth rings distinct to faint, with wavy ma pits similar to intervessel pits but half­ boundaries, marked by rows of weakly radi­ bordered. Helical thickenings fine and close­ ally flattened latewood fibres. Vessels dif­ ly spaced, throughout body of some vessel fuse, 136-138/sq.mm, 61-75% solitary, elements. Gummy contents absent, remainder in tangential and oblique multiples Fibre-traeheids 900 (620-1130) um long, of 2-4, round to oval, tangential diameter F/V ratio 1.7, medium thick- to very thick- 31-35 (15-58) um, radial diameter 40 (17-

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70) um, walls 1-2 um thick. Vessel element Rosa L. (Figs. 6, 19,35; Tab1e 14) length 380-440 (180-570) um, L/D ratio Material studied: R. cymosaTratt.: Huo­ 12-13. Perforations almost exclusively sim­ shan, Anhui, He Yunhe s.n ., diam. 20 rnm, ple in oblique end walls , but sporadic irregu­ alt. 150 m. - R. henryi Bou1eng.: Huoshan, lar multiple perforations also found in one Anhui, He Yunhe s.n., diam. 19 rnrn, alt. sample (Huang Chenglin s.n.). Intervessel 200 m; Fujian, CAFw19794, diarn . 17 mm. pits nonvestured, alternate and scalarifonn to -R. Iaevigata Michx.: Hefei, Anhui, S.Y . oppo site, elongate to oval, 3-20 umin diam­ Zhang s. n., cu1t., diam. 16 mm, alt. 50 m; eter, with slit-like apertures. Vessel-ray and Fujian, CAFw 19798, diam. 10 mm.- R. vessel-parenchyma pits with reduced bor­ macrophylla Lindl.: Tibet, CAFw 19331 , ders, alternate to scalarifonn. Helical thicken­ diam. 35 mm. - R. multiflora Thunb.: Hu­ ings absent. Gummy contents absent. angshan, Anhui, Liu Xiumei s.n ., diam . 15 Fibre-tracheids 710-1100 (530-1520) mm , alt. 350 m. - R. roxburghii Tratt .: Tun­ um long, F/V ratio 1.9-2.4, medium thick­ xi, Anhui, Ye Shuyou s.n ., diarn. 5 mm, alt. to very thick-walled, with distinctly bordered 200 m; Huangshan, Anhui, Liu Xiumei s.n ., pits of 4-5 um in diameter, densely placed diam. 10 mm, alt. 400 m. - R. sertata Rolfe : in radial and tangential walls. Helical thicken­ Hefei, Anhui, S.Y. Zhang s.n ., cult. , diam. ings absent in one sample (Huang Chenglin 19 mm, alt. 50 m. - R. xanthina Lindl .: She­ s. n.), but conspicuous in another sample, nyang, Liaoning, Qian Hong s.n ., diam . 11 present in almost all fibre-tracheids. mm. Parenchyma sparse to fairly common, Deciduous, erect, scrambling or climbing predominantly apotracheally diffuse, rarely shrubs distributed throughout China, but scanty paratracheal, in 2-3 (l-5)~celled mainly from Northwest and Southwest Chi­ strands. na, widely cultivated as ornamentals. Rays 12-13 (7-19)/mm, of two distinct Growth rings distinct, with more or less sizes : narrow and low rays 1(-2)-seriate, and wavy boundaries, marked by differences in wide and tall rays (3-)4-7-seriate. Uniseri­ vessel diameter between latewood and sub­ ate rays 7-12 (2-26) cells high, composed sequent earlywood, locally inflated broad exclusively of upright cells; multiseriate rays rays, and usually also by 1-4 rows of radial­ 7.4 -18.0 (0.3-27.5) mm high , composed 1y flattened latewood fibres . Wood ring-por­ of square (to weak1y procumbent) body cells, ous, with 1-2 rows of large earlywood ves­ upright marginal cells and weakly differen­ sels. Latewood vessels 114-343 (mostly tiated sheath cells. 114-176)/sq. mm , 40-96% solitary (all Crystals sparse or common, prismatic, above 60% except in R. macrophylla), re­ medium-sized, present in non-en1arged ray mainder in oblique and tangential multiples cells. Pith flecks and traumatic gum ducts ab­ of 2-5, round to oval or angular, tangential sent. diameter of earlywood vessels 35-150 (30­ Notes: 1. Metcalfe and Chalk (1950) men­ 220) um, radial diameter 45-210 (40-250) tioned that uniseriate rays in Kerria are ab­ um; tangential diameter of latewood vesse ls sent. However, they are common in the ma­ 19-43 (15-i15) um, radial diameter 23-51 terial we examined. In addition, ray frequen­ (18-125) um, walls 1-3 um thick. Vessel cy in the two sampies of the species is higher element length 280-400 (210-930) um . than 3-5/mm reported by them. Perforations exclusively simple in oblique to 2. Solereder (1908) and Greguss (1959) almost horizontal end walls . Intervessel pits recorded occasional fibres with simple pits in alternate, round, oval to polygonal, 3-10 um Kerria, However, ground tissue in the speci­ in diameter, with slit-like apertures, aperture s mens we examined are composed exclusivel y occasionally coalescent in some species. Ves­ of fibre-tracheids with distinctly bordered sel-ray and vessel-parenchyma pits simi1ar pits . to intervessel pits but half-bordered and slight­ 3. Occasional scalarifonn intervessel pits Iy smaller, sometimes with slightly reduced were also observed by Cutier et al. (1987) in borders, 2-7 um in diameter. Helic a1 thick­ rootwood of Kerriajaponica . enings fine, discontinuous and irregu1arly

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Table 14. Variation in selected wood anatomical features of Rosa.

1 2 3 4 5 6 7 8 9 10 11 R. cymosa 172 63 43 94 360 5-8 660 1.9 15 14.3 R R. henryi 160 64 32 153 390 5-10 970 2.5 16 5.0 R R. laevigata Zhang s.n, 122 91 34 71 340 5-7 650 1.9 12 3.0 R CAFw 19798 114 96 32 87 400 5-7 870 2.2 7 4.6 R R. macrophylla 176 40 30 71 310 5-7 860 2.8 16 1.5 R. multiflora 123 65 33 101 310 3-8 740 2.4 12 10.0 R R. roxburghii 140 88 28 60 360 4-9 630 1.7 5 5.9 R R. sertata 150 84 27 93 280 5-7 620 2.2 11 2.9 R. xanthina 343 92 19 35 310 4-7 510 1.7 5 1.0 1: Vessel frequency (/sq.mm); 2: Pereentage of solitary vessels; 3: Average tangential diameter of latewood vessels (J.lID); 4: Average tangential diameter ofearlywood vessels (J.lID); 5: Average vessel element length (J.lID); 6: Size of intervesseI pits (J.lID); 7: Average fibre-tracheid length (J.lID); 8: FIV ratio; 9: Width of the widest rays (number of cells); 10: Average multiseriate ray height (mm); 11: Prismatic crysals present in ray cells (R) or absent (-).

spaced, mainly presentin body of narrow Crystals sparse to common, prismatic, oc­ vessel elements. Gummy contents absent. casionally elongated, medium-sized, present Vasicentric tracheids present in a few sam­ in non-enlarged ray cells, crystals not ob­ ples, but only weakly differentiated from served in R. macrophylla, R. sertata and R. ground tissue, mainly confined to earlywood, xanthina. Pith flecks usually absent, but in­ generally shorter than ground tissue fibres frequently present in R. cymosa. Traumatic but larger in diameter, with distinctly border­ gum ducts absent. ed pits of 4-8 um in diameter and irregular Notes: 1. Sporadic multiple perforations in shape. were reported for two Himalayan species by Fibre-tracheids 510-970 (350-1200) J.lID Suzuki et al. (1991). long, F/V ratio 1.7-2.8, medium thick- ro 2. Helical thickenings in many fibres of very thick-walled, with distinctly bordered R. arabica and R. pulverulenta were observed pits of 3-7 J.lID in diameter, in radial and tan­ by Fahn et al. (1986). However, helical gential walls. Helical thickenings occasion­ thickenings in fibres are mostly absent in the ally present in a few samples, very fine. Chinese species. Parenchyma usually scanty (but abundant 3. Ray frequency (3-5/mm) in the genus in R. macrophylla), apotracheally diffuse and reported by Metcalfe and Chalk (1950) is scanty paratracheal, in 2-5(-8)-eelled strands. much lower than in our material . Rays 6-15 (4-20)/mm, of two distinct 4. Diffuse-porous rootwood of Rosa was sizes : narrow and low rays 1(-2)-seriate, and described by Cutler et al. (1987) . Ring-por­ wide and tall rays (3-)4-16-seriate (the ous stern wood in the genus was also report­ widest rays 5-seriate in R. roxburghii and R. ed by Greguss (1959), Grosser (1977) and xanthina to Iö-seriate in R. henryi and R. Schweingruber (1990). macrophylla). Uniseriate rays 4-10 (1-39) cells high, composed exclusively of upright cells; multiseriate rays 1.0-14.3 (0.2-22.5) Rubus L. (Figs. 1,53; Table 15) mrn high, usually composed of procumbent Material studied: R. biflorus Buch-Harn, to square body cells, upright marginal cells ex Srnith: Tibet, CAFw 19326, diam. 3 cm.­ and weakly differentiated sheath cells. R. chingii Ru: Qingliangfeng, Anhui, Huang

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Table 15. Variation in selected wood anatomical features ofRubus.

2 3 4 5 6 7 8 9 10 11 R. biflorus 97 43 48 300 - 5-14 710 2.4 10 4.2 R. chingii 90 50 55 420 + 4-10 790 1.9 9 9.4 R. coreanus 102 71 49 440 5-6 800 1.6 10 10.0 R. corchorifolius 85 69 37 460 4-9 740 1.8 10 3.6 R. swinhoei 128 56 36 670 4-5 890 1.3 9 15.0 R. trianthus 84 46 46 420 4-7 790 1.9 9 14.1 R

1: Vessel frequency (lsq.mm); 2: Percentage of solitary vessels ; 3: Average tangential vessel diameter (um); 4: Average vessel element length (11m); 5: Irregular perforations present (+) or absent (-); 6: Sizes of intervessel pits (um); 7: Average fibre-tracheid length (um); 8: F/V ratio; 9: Width of the widest rays (number of cells); 10: Average multiseriate ray height (mm); 11: Prismatic crystals present in ray cells (R) or absent (-).

Chenglin s.n., diam . 18 mm, alt. 350 m. ­ tangential diameter of large vessels 84 (45­ R. coreanus Miq.: Qingliangfeng, Anhui, Hu­ 113) um, radial diameter 104 (55-133) 11m. ang Chenglin s. n., diam . 10 mm, alt. 400 m. Vessel element length 300-670 (130-1220) -R. corchorifolius L. f.: Fujian , CAFw 11m. L/D ratio 6-12(-19). Perforations ex­ 20058, diam. 20 mm. - R. swinhoei Hance : clusively simple in oblique to almost horizon­ Fujian, CAFw 20057 , diam. 15 mm. - R. tal end walls, but sporadic irregular perfora­ trianthus Focke : Tunxi, Anhui , Ye Shuyou tions also found in first formed secondary s.n. , diam. 15 mm, alt. 200 m. xylem of R. chingii. Intervessel pits nonves­ Deciduous , rarely evergreen, shrubs, sub­ tured, alternate, polygonal, oval to round, shrubs, scrambling shrubs or herbs distribut­ 4-10(-14) 11m in diameter, with slit-like ed throughout China, but mainly from South­ apertures. Vessel-ray and vessel-parenchy­ west China, This large genus is represented ma pits similar to intervessel pits but half­ by almost 200 species in China. bordered and slightly smaller, or with some­ Growth rings faint to absent in most sam­ what reduced borders. Helical thickenings pies studied. Wood semi-ring-porous to dif­ absent. Gummy contents absent. fuse-porous. The vessel diameter usually de­ Fibre-tracheids 710-890 (430-1340) 11m crease s gradually from pith outward, 84­ long, F/V ratio 1.3-2.4, medium thick- to 128/sq.mm, 43-71% solitary, remainder very thick-walled, with distinctly bordered predominantly in tangential, rarely oblique pits of 3-4 um in diameter, in radial (densely and radial multiples of 2-5, oval to round or placed) and tangential walls. Helical thicken­ weakly angular, tangential diameter 36-55 ings absent. (17-110) 11m, radial diameter 43-69 (18­ Parenchyma sparse, apotracheally diffuse 155) 11m, walls I-211m thick. Vessels of and scanty paratracheal, in 3-4 (2-7)-celled two distinct sizes in R. biflorusand R. trian­ strands , thus, small vessels randomly scattered in R. Rays 9-13 (4-17)/mm, of two distinct biflorus, tangential diameter of small vessels sizes: narrow and low rays 1(-2)-seriate, 40 (26- 48) 11m, radial diameter 52 (30- 60) and wide and tall rays 3-9(-1O)-seriate (the 11m; tangential diameter of large vessels 95 widest rays 9-seriate in R. chingii, R. swin­ ( 80- 1 1 5) ~ , radial diameter 132 (120- 155) hoei and R. trianthus to lü-seriate in R. bi­ 11m; but small vessels in R. trianthus OCCUT­ florus, R. coreanus and R. corchorifolius). ring between two wide rays from pith out­ Uniseriate rays 7-16 (2- 61) cells high, com­ ward, tangential diameter of small vessels 40 posed exclusively of upright cells; multiseri­ (18-50) 11m, radial diameter 47 (20- 68) 11m; ate rays 3.6-15.0 (0.5- 45.0) mm high,

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 69 composed of upright and square cells and clusively simple in oblique end walls. Inter­ weakly differentiated sheath cells. vessel pits nonvestured, alternate, polygonal Crystals mostly absent, but infrequent in or round to oval, 3-9 11m in diameter, with 1<. trianthus, prismatic, small to medium­ slit-like apertures, apertures occasionally co­ sized, usually one crystal (occasionally two) alescent in P. sinensis. Vessel-ray and ves per non-enlarged ray cell. Pith flecks and sel-parenchyma pits with reduced borders or traumatic gum ducts absent. fully half-bordered and slightly smaller than Notes: 1. The drawings by Greguss (1959) intervessel pits, 3-6 11m in diameter. Helical indicate incidence of libriform fibresin Rubus. thickenings fine in P. sinensis or coarse in We did not observe libriform fibres in six P. utilis, usually closely spaced, throughout Chinese species studied here. body of most vessel elements. Gummy con­ 2. Moderately large vessels (200-300 11m tents present in some vessels. in diameter) recorded for Rubus by Metcalfe Fibre-tracheids 700-780 (490-980) 11m and Chalk (1950) were not observed in the long, F/V ratio 1.9-2.2, very thick-walled, Chinese species. with distinctly bordered pits of 4-7 11m in 3. Ray frequency (3-5/mm) in the genus diameter, in radial and tangential walls. Heli­ documented by Metcalfe and Chalk (1950) is cal thickenings fine, present in a few fibre­ much lower than in our material. tracheids in P. sinensis, or coarse and con­ 4. Indistinct growth rings were also report­ spicuous, in almost all fibre-tracheids in P. ed by Schweingruber (1990), and were also utilis. described by Cutler et al. (1987) in rootwood Parenchyma scanty (P. sinensis) to moder­ of the genus. ately common (P. utilis), predominantly apo­ tracheally diffuse, but diffuse-in-aggregates Subfamily IV: Prunoideae Focke and scanty paratracheal also present in P. uti­ lis, in 3-4 (2-7)-celled strands. Amygdalus L., see Prunus s.l. Rays 13 (9-17)/mm, 1- 2(- 3)-seriate in Armeniaca Mill., see Prunus s.l. P. sinensis up to 4(-5)-seriate in P. utilis. Cerasus Mill., see Prunus s.l. Uniseriate rays 2-6 (1-16) cells high, com­ Laurocerasus Tourn. ex Duh., see Prunus posed of square and upright cells; multiseriate s. l. rays 190-550 (100-1757) 11m high, Padus Mill., see Prunus s.l. composed of procumbent (occasionally to Pygeum Gaertn., see Prunus s.l. square) body ray cells and 1-5 rows of square to upright marginal cells. Prinsepia Royle Crystals absent in P. sinensis or sporadi­ Material studied: P. sinensis (Oliv.) Oliv. cally present in P. utilis, prismatic, medium­ ex Bean: Fusuen, Liaoning, HEFw 1582, sized, in non-enlarged ray cells. Pith flecks diam. 20 mm, -Poutilis Royle: Tibet, CAFw and traumatic gum ducts absent. 19341. Notes : 1. Predominantly solitary vessels Deciduous erect or climbing shrubs from and semi-ring-porous wood were also report­ northern regions of China. ed by Kanehira (1921a) for Prinsepia scan­ Growth rings distinct, marked by rows of dens. radially flattened latewood fibres and differ­ 2. The two Chinese species differ from ences in vessel diameter between latewood each other in certain features (e.g. helical and subsequent earlywood. Wood semi-ring­ thickenings in vessel elements and fibre-tra­ porous to diffuse-porous , Vessels 148-313/ cheids, axial parenchyma, ray size, and crys­ sq.mm, 68-87% solitary, remainder mainly tals). in oblique, rarely radial and tangential multi­ 3. Although Prinsepia shares some wood ples of 2-3, angular or oval to round, tan­ anatomical features with Prunus s.l. in the gential diameter 25-30 (15-65) 11m, radial sense of Kalkrnan (mainly parenchyma fea­ diameter 31-44 (20-95) 11m, walls I-211m tures and crystal features), it would be some­ thick. Vessel element length 350-370 (230­ what aberrant in the Prunus s.l. alliance. 620) 11m. LID ratio 12-15. Perforations ex- Helical thickenings are present in the fibre-

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Table 16. Variation in selected wood anatomical features ofAmygdalus.

2 3 4 5 6 7 8 9 10 11 12 13

A. davidiana 144 26 30 65 270 770 2.7 1-3 4 0.8 D - A. kansuensis 52 68 37 84 320 - 1140 3.6 2-5 4(-5) 0.5 D - A. mira 159 38 48 109 310 - 1040 3.3 3-4 5 0.7 D - A. persica HEFw 3803 344 28 31 82 270 820 3.0 3-4 5 0.6 - + CAFw 15876 52 44 36 68 280 910 3.3 2-5 5(-6) 0.4 Huang s.n . 181 28 40 69 290 + 580 2.0 1-4 3(-4) 0.7 D + CAFw 5716 192 54 29 76 280 770 2.8 2-5 5(-6) 0.7 HEFw 345 166 25 35 86 330 900 2.7 2- 5 4(-5) 0.5 - + CAFw16900 107 30 45 74 390 + 960 2.5 1-4 5(-6) 0.5 A. triloba CAFw 13721 330 32 38 60 330 710 2.2 4-5 3(- 4) 0.4 D - Qian Hong s.n. 600 47 23 40 230 690 3.0 3-5 4 0.5 - +

1: Vessel frequency (/sq.mm); 2: Percentage of solitary vessels; 3: Average tangential diameter oflatewood vessels (um); 4: Average tangential diameter of earlywood vessels (um); 5: Average vessel element length (um); 6: Multiple perforations present (+) or absent (-); 7: Average fibre length (um); 8: F/V ratio; 9: Size of pits in fibres (um) ; 10: Width of the widest rays (number ofcells); 11: Average multiseriate ray height (mm); 12: Druses (D) present in ray cells or absent (-); 13: Traumatic gum ducts present (+) or absent (-).

tracheids ofPrinsepia, while they are usually alt. 250 m; Guangxi, CAFw 15876; Qing­ absent in Prunus s.l.; ray size and composi­ liangfeng, Anhui , Huang Chenglin s.n., tion are also different from most Prunus s.l. diam. 15 mm, alt. 300 m; Qingtao, Shan ­ species. In addition, the genus has a higher dong, CAFw 5716; Huoshan , Anhui , HEFw percentage of solitary vessels (larger than 345, alt. 250 m; Guangdong, CAFw 16900. 60%) and higher LID ratio than Prunus s.l. - A. triloba (Lindl.) Ricker: Beijing, CAFw 1372 1, diam. 30 mm; Shenyang, Liaoning, Prunus s.l. (cf. Kalkman 1965)1 Qian Hong s.n., diam. 15 mm. Deciduous trees or shrubs mainly distri­ Amygdalus L. (Figs. 21,24; Table 16) buted in western regions and Northwest Chi­ Material studied: A. davidiana (Carr.) C. na, widely cultivated throughout the country de Vos ex Henry: Shenyang, Liaoning, Qian as fruit trees. Hong s.n. , diam. 25 mm. - A. kansue nsis Growth rings distinct, marked by differ­ (Rehd.) Skeel: Tibet, CAFw 19342. -A. mira ences in vessel diameter between latewood (Koehne) Yu & Lu: Tibet, CAFw 19344. ­ and subsequent earlywood and rows of A. persica L.: Dongzhi, Anhui, HEFw 3083, weakly radially flattened latewood fibres. Wood ring-porous (rarely ranging from ring­ 1) The concept of Prunus s.l. or the Prunus porous to semi-ring-porous), with 1-2(-4) alliance here follows Kalkman (1965) and rows of large earlywood vessel s. Latewood includes Amygdalus, Armeniaca, Cerasus, vessels 52-344I sq. mm (mostly above Laurocerasus, Padus, Prunus s.str. and lOO/sq.mm), 25-68% solitary (all below Pygeum. This broad concept differs slight­ 54%, except A. kansuensis), remainder in ly from Bentham and Hooker's delimi­ radial, oblique and tangential multiples of tation (1865) who treated Pygeum as a 2-7, oval to round, tangential diameter of separate genus. earlywood vessels 60-109 (20-150) um,

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 71 radial diameter 88-120 (26-190) 11m; tan­ cells ofA. davidiana, A. kansuensis , A. mira gential diameter of 1atewood vesse1s 29-48 andA. triloba (CAFw 13721), medium-sized, (12-70) 11m, radial diameter 35-53 (18­ one crystal per non-enlarged ray cell, Pith 97) 11m, walls 1-3lJ.ffi thick. Vesse1 element flecks usually absent, but infrequent in some 1ength 270-390 (140- 970) 11m. Perfora­ specimens of A.persica. Traumatic gum ducts tions exc1usive1y simple in oblique (late­ present in A. persica (HEFw 3803, HEFw wood) to almost horizontal (ear1ywood) end 345 and Huang Chenglin s. n.) and A. triloba walls, but sporadic scalariform and multiple (Qian Hong s.n .), usually in short to long perforations with 3-5 bars noted in A. persi­ tangential bands. ca (CAFw 16900 and Huang Chenglin s.n.) . Notes: 1. Ring-porous, ring-porous to Intervesse1 pits nonvestured, alternate, poly­ semi-ring-porous, and diffuse-porous wood gonal, round to oval, (3-)4-9 11m in diarn­ in A. communis was reported by Fahn et al. eter, with slit-like apertures, apertures rarely (1986) , Oprea (1972), and Jacquiot et al. coalescent in A. davidiana, A. kansuenesis (1973), respectively. All Chinese species ex­ and A. persica (HEFw 345). Vessel-ray and arnined are characterised by ring-porous wood vessel-parenchyma pits c1ear1y smaller than (rarely ring- to semi-ring-porous). Ring-por­ intervesse1 pits and half-bordered with re­ ous wood in the genus was also reported by duced borders, 2-5 11m in diameter. Helical Greguss (1959) and Schweingruber (1990) . thickenings well developed, usually wide1y 2. Opposite intervessel pits in A. commu­ spaced, throughout body of vessel elements. nis reported by Jacquiot et al. (1973) were Gummy contents present in some vessels of not observed in the species we studied. all species. 3. Fahn et al. (1986) noted tyloses in A. Vasicentric tracheids sparse in some sam­ communis. However, no tyloses were noted ples, main1y in earlywood, with distinctly in our material. bordered pits in both radial and tangential 4. In the generic description quantitative walls and irregular in shape. data for the immature sample (Qian Hong) of Fibres 770-1140 (480-1400) 11m long, A. triloba have been left out due to its signifi­ FIV ratio (2.2-)2.7-3.6, very thin- to very cant differences from all the other samples. thick-walled, with (minutely to) distinctly bordered pits of (2-)3-5 11m in diameter in Armeniaca Mill. (Figs. 9, 28, 37, 61; Table A. kansuensis, A. mira and A. triloba, large­ 17) 1yrestricted to the radial walls, but in A. da­ Material studied: A. holosericea (Batal.) vidiana and some samples of A. persica, Kost.: Zhongtiaoshan, Shanxi, HEFw 131.­ fibres with simple to minutely bordered pits A. mandshurica (Maxirn.) Skv. : Heilong­ of I-311m in diameter also noted. Helical jiang, Northeast Forestry University s.n. ­ thickenings absent. A. mume (Sieb. & Zucc.) Carr.: Hefei , An­ Parenchyma usually scanty, apotracheally hui, S.Y. Zhang s.n ., diam. 25 mm, alt. 50 diffuse and scanty paratracheal, in 2-4 (2-7)­ m; Tibet, CAFw 19345; Guangdong, Gui­ celled strands. zhou Forest Institute s. n. - A. mume var. Rays 8-14 (6-18)/mm, of two sizes: alphandii Carr.: location unknown: CAFw narrow and low rays 1(-2)-seriate, and wide 7700. -A. sibirica CL.) Lam.: Zhongtiaoshan, and tall rays (2-)4-5(-6)-seriate, but in Shanxi, HEF 134. -A. vulgaris Lam.: Xing­ four juvenile samples , rays 1-3(-4)-seriate. jiang, CAFw 13823; Wuhu, Anhui, HEFw Uniseriate rays 3-10 (1-36) cells high, com­ 716, alt. 150 m; Xinjiang, HEFw 2653; Tai­ posed of square and upright cells, multiseri­ baishan, Shanxi, CAFw 588; Zhongtiaoshan, ate rays 430-840 (30-2000) 11m high, com­ Shanxi, HEFw 125; Shenyany, Liaoning, posed of procumbent (to square in juvenile Qian Hong s.n., diam. 10 mm. samples) body cells and 1-4(-7) rows of Deciduous trees, rarely shrubs mainly dis­ square (to upright) marginal cells. tributed in the northem regions of Qinling Crystals absent in A. persica except in one and Huaihe , especially in the Yellow River sample (Huang s.n .) and A. triloba (Qian valley, widely cultivated as fruit trees and or­ Hong s.n .), but sporadic druses noted in ray namentals.

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Table 17. Variation in selected wood anatomical features of Armeniaca.

2 3 4 5 6 7 8 9 10 11 12 13

A. h%sericea 288 31 30 79 260 4-6 490 1.9 6(-8) 0.6 D-R + R A. mandshurica 162 36 38 80 290 5-8 760 2.6 6(-8) 0.6 D-R + R A. mume Zhang s.n. 335 49 30 230 4-6 750 3.2 5 0.5 + RtoS CAFw 19345 233 37 38 310 6-9 780 2.5 4(-5) 0.7 R Guizhou s,n. 110 52 36 290 4-6 840 2.9 8 0.6 + StoD var. a/phandii 169 50 34 280 4-8 740 2.7 6(-8) 0.9 D-A - StoD A. sibirica 175 30 32 290 4-6 780 2.7 6(-8) 0.4 D-R + A. vu/garis CAFw 13823 204 25 39 78 280 5-9 770 2.8 6(-7) 0.7 D-R -R HEFw 716 325 48 30 73 290 4-7 720 2.5 8 0.6 + R HEFw 2653 287 15 37 91 240 5-7 490 2.0 6(-7) 0.6 D-R -R CAFw 588 286 32 37 90 240 5-7 490 2.1 6(-8) 0.5 D-R -R HEFw 125 279 31 35 104 260 5-8 570 2.2 6(-7) 0.5 D-RA -R Qian Hong s.n.340 48 24 60 250 3-7 550 2.2 4(-5) 0.6 D-R -R

1: Vessel frequency (/sq.mm); 2: Percernage of solitary vessels; 3: Average tangential diameter of latewood vessels (1Jll1); 4: Average tangential diameter of earlywood vessels (1Jll1); 5: Average vessel element length (um); 6: Size of intervessel pits (1Jll1); 7: Average fibre length (1Jll1); 8: F/V ratio; 9: Width of the widest rays (number of cells); 10: Average multiseriate ray height (mm); 11: Druses (D) present in ray cells (R), axial parenchyma cells (A), in both (RA) or absent (-); 12: Traumatic gum ducts present (+) or absent (-); 13: Wood ring-porous (R), semi-ring-porous (S) or diffuse-porous (D).

Growth rings distinct, marked by differ­ 4-9 um in diameter, with slit-like apertures, ences in vessel diameter between latewood apertures occasionally coalescent in A. vulga­ and subsequent earlywood and rows of radi­ ris (Qian Hong s.n.), Vessel-ray and vessel­ ally flattened latewood fibres. Wood ring­ parenchyma pits clearly smaller than interves­ porous (except in A. mume which varies sei pits and half-bordered with reduced bor­ from ring-porous to diffuse-porous), with ders, 2-5 um in diameter. Helical thicken­ 1-2(-3) rows of large earlywood vessels. ings usually weIl developed, widely spaced, Latewood vessels 1l0-340/sq.mm (mostly throughout body of vessel elements. Gummy 200-340/sq.mm), 15-52% solitary, re­ contents present in most species, but absent mainder in oblique, radial and tangential mul­ in A. mume (Zhang s.n. and Guizhou s.n.). tiples of 2-5, oval to round or weakly angu­ Vasicentrlc tracheids sparse in a few sam­ lar, tangential diameter of earlywood vessels ples, mainly in earlywood, with distinctly 60-104 (35-160) um, radial diameter 72­ bordered pits in both radial and tangential 126 (46-180) um; tangential diameter of walls, usually irregular in shape. latewood vessels 24-39 (8-60) um, radial Fibres 490-840 (280-1310) um long, diameter 29-53 (12-127) um, walls 1-2 FIV ratio 1.9-3.2, medium thick- to very 11m thick. Vessel element length 230-310 thick-walled , with distinctly bordered pits of (140-540) um. Perforations exclusively (2-)3-5 um in diameter, largely restricted to simple, usually in oblique (latewood vessels) radial walls. Helical thickenings absent. to almost horizontal (some earlywood ves­ Parenchyma usually scanty, apotracheally sels) end walls. Intervessel pits nonvestured, diffuse and scanty paratracheal, in 3-4 (2-6)­ alternate, polygonal or oval to round, (3-) celled strands.

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Rays 7-12 (5-14)/mm, oftwo sizes: nar­ 2. Vasicentric traeheids and traumatic gum row and low rays 1(-2)-seriate, and wide duets in A. vulgaris were also reported by and tall rays (2-)4-6(-8)-seriate. Uniseriate Fahn eraJ. (1986). rays 4-9 (1-25) cells high, composed of 3. In Prunuss.l. (cf. Kalkman 1965), on­ square to weakly procumbent, and/or up­ ly Amygdalus and Armeniaca species are char­ right cells; multiseriate rays 450-880 (100­ aeterised by ring-porous wood, whereas the 1650) 11m high, composed of procumbent remaining species are usually diffuse-porous, body cells and 1-5 rows of square to upright rarely ranging to semi-ring-porous, Further­ marginal cells (heterogeneous m to Il), but more, Amygdalus and Armeniaca are very in rwo shrub specimens of A. mume and A. similar to each other in wood structure and vu/garis (Qian Hong s.n.), multiseriate rays cannot be differentiated wood anatomically. composed of procumbent to square body 4. Greguss (1959) noted homogeneous cells and several rows of square (to upright) rays in Prunus armeniaca which is different marginal cells. from our results. Crystals sparse to frequent, druses, me­ dium-sized, in non-eharnbered, normal or Cerasus Mill. (Figs. 34, 46, 54, 56, 58; Table weakly enlarged ray cells; occasionally in 18) chambered, non-enlarged axial parenchyma Material studied: C. avium (L.) Moench in A. mume var. alphandii and A. vu/garis var. decumana Mordant: Guangxi, HEFw (HEFw 125), one crystal per cell or chamber; 1096. - C. campanulata (Maxim.) Yu & Li: chains of up to 10 charnbers in axial paren­ Guaiigdong, South China Agrlcultural Uni­ chyma cells ofA. mume var. alphandii, crys­ versity 2333; Fujian, CAFw 13055; Taiwan, tals not observed in A. mume and one sarnple CAFw 5040; Northeast China, Northeast of A. vu/garis (HEFw 716) . Pith flecks rare Forestry University s. n. - C. cerasoides to common in A. ho/osericea,A mandshurica (D. Don) Sok.: Guangdong, FRIGw 1208; andA. vu/garis (HEFw 125 and CAFw 588). Yunnan, Yunnan Academy of Forestry 363. Traumatic gum ducts present in A. holoseri­ -C. c/arofo/ia (Schneid.) Yu & Li: Zhong­ cea, A. mandshurica, A. mume (Zhang s.n . tiaoshan, Shanxi, HEFw 126; Yunche, Yun­ and Guizhou Forest Institute s.n.), A. sibirica nan, Yunnan Academy of Forestry 368, alt. and A. vu/garis(HEFw 716), in long tangen­ 1800 m; Zhongtiaoshan, Shanxi, HEFw tial bands. 129. - C. conradinae (Koehne) Yu & Li: Dali, Notes: 1. Ring-porous wood or ring-por­ Yunnan, Yunnan Academy of Forestry 205, ous to semi-ring-porous wood in Prunusar­ alt. 2200 m. - C. die/siana (Schneid.) Yu & meniaca (Armeniaca vu/garis) was also re­ Li: Jiangxi, CAFw 17429; Taiping, Anhui, ported by Greguss (1959), Schweingruber HEFw 3884, alt. 500 m. - C. maximowiczii (1990), Fahn era/. (1986) and Jacquiot era/. (Rupr.) Kom.: Changbaishan, Jining, Qian (1973). All Chinese species studied are char­ Hong s.n., diarn. 16 mm. - C. pseudocera­ acterised by ring-porous wood except Arme­ sus (Lindl.) G. Don: Anhui, HEFw 550; niaca mume (or Prunus mume) where the Guizhou, Guizhou Forestry Institute s.n.; wood varies from ring-porous, via semi-ring­ Zhongtiaoshan, Shanxi, HEFw 130; Guang­ porous to diffuse-porous. The variation of dong, FRIGw 463; Guangdong, CAFw porosity in the species may be related to hor­ 15053. - C. serru/a (Franch.) Yu & Li: Sich­ ticultural treatment. Armeniaca mume origi­ uan, CAFw 46; Huangshan, Anhui, Liu Xiu­ nates from China and has been cultivated mei s.n., diarn. 23 mm, alt. 600 m. - C. ser­ widely as an important ornamental in China rulata(Lindl.) G. Don ex Loud.: Jixi, Anhui, for over 3000 years (Yu 1986). Nowadays HEFw 3160, alt. 500 m. - C. serrulata var. many varieties (cf. Yu 1986) are developed sachalinensis Fr. Schmidt: location unknown, by hybridisation, grafting and modem tech­ CAFw 4199. - C. setu/osa(Batal.) Yu & Li: niques. Semi-ring-porouswood in both nor­ Zhongtiaoshan, Shanxi, HEFw 133; Taibai­ mal and dwarf Prunusmume (or Armeniaca shan, Shanxi, CAFw 564. - C. szechuanica mume) was also reported by Baas er a/. Yu & Li: Tibet, CAFw 19339. - C. tomen­ (1984). rosa (Thunb.) Wall.: Tibet, CAFw 19337,

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Table 18. Variation in selected wood anatomical features of Cerasus.

2 3 4 5 6 7 8 9 10 11 12 C. avium var. decumana 220 22 50 390 6-10 1000 2.6 3-5 4(-5) 0.7 PD-AR - C. campanulata CAFw 13055 91 20 59 390 5-7 1000 2.4 3-5 4(-5) 0.8 P-A CAFw5040 109 27 60 370 5-8 1080 2.9 2-4 4(-5) 0.5 P-R 2333 63 29 53 440 5-7 1400 3.2 3-4 5(-6) 0.7 P-A Northeast s.n. 105 28 44 400 5-7 1200 3.0 3-5 3 0.6 P-A C. cerasoides FRIGw 1208 85 38 50 370 4-6 1070 2.9 3-4 5(-6) 0.8 P-A + 363 86 28 64 390 4-6 1120 2.8 4-5 6(-8) 0.7 P-AR - C. clarofolia HEFw 126 94 27 45 380 5-8 1230 3.2 4-5 4(-5) 0.3 D-A 368 88 18 65 400 4-8 1160 2.9 4-5 6(-7) 1.5 P-AR - HEFw 129 162 25 37 340 5-8 940 2.8 3-4 4(-5) 0.4 D-R - C. conradinae 56 11 65 400 5-10 1370 3.4 3-4 5(-6) 0.5 C. dielsiana CAFw 17429 125 32 42 500 5-10 940 1.9 3-5 4 0.6 P-A HEFw 3884 185 22 42 350 6-8 980 2.8 3-5 5(-6) 0.6 P-A C. maximowiczii 283 33 28 290 4-7 600 2.0 1-4 2(-3) 0.4 D-A - C. pseudocerasus HEFw 550 85 36 45 5-9 2-4 4(-5) 0.5 P-A Guizhou s.n. 136 26 53 480 7-11 900 1.9 3-4 4(-5) 0.6 P-A HEFw 130 119 43 38 300 5-8 870 2.9 3-5 5(-6) 0.5 D-A C. serrula CAFw46 58 43 55 360 7-11 770 2.2 2-4 4(-5) 0.6 PD-AR - Liu Xiume i s.n. 186 32 27 320 3-6 940 2.9 3-4 4(-5) 0.8 D-A - C. serrulata 192 45 36 270 4-5 740 2.8 4-5 5(-7) 0.6 D-A var. sachalinensis 103 20 50 390 5-8 1020 2.6 3-4 4 0.4 PD-AR - C. setu/osa HEFw 133 117 19 47 460 5-10 930 2.0 2-4 4 0.4 D-A - CAFw564 53 54 40 370 5-7 1010 2.7 2-4 4 0.4 D-A - C. tomentosa 360 20 23 300 3-6 560 1.9 3-5 3 0.5 C. yedoensis 115 50 35 360 4-8 770 2.2 1-4 4 0.4 DP-A +

1: Vessel frequency (/sq .mm); 2: Percentage of solitary vessels; 3: Average tangential vessel diameter (um); 4: Average vessel element length (um); 5: Size of intervessel pits (um); 6: Aver- age fibre length (um); 7: F/V ratio; 8: Size of pits in fibres (um); 9: Width of the widest rays (number of ceIls); 10: Average multiseriate ray height (mm); 11: Prismatic crystals (P), druses (0), or both (PD) present in axial parenchyma (A), rays (R), in both (AR) or absent (-); 13: Traumatic gum ducts present (+) or absent (-) . diam. 25 mm. - C. yedoensis (Matsum.) Yu Growth rings distinct, marked by rows of & Li: Hefei, Anhui, S.Y. Zhang s.n., cult., radially flattened latewood fibres, in some diam. 20 mm, alt. 50 m. species by weak differences in vessel diam- Deciduous trees or shrubs, most from eter between latewood and subsequent early- western regions and Southwest China, wide- wood as weIl. Wood mostly diffuse-porous, ly cultivated as ornamentals and fruit trees. sometimes ranging to semi-ring-porous. Ves-

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sels in a weakly oblique pattern in C. campa­ ing some procumbent central cells; multiseri­ nulata (CAFw 13055 and Northeast Forestry ate rays 320-1490 (100-3100) 11m high, University s.n.) and C. claroiolia (HEFw composed of procumbent body cells and 1-4 126), 53-360/sq.mm (mostly 80-220/ (-5) rows of square (to upright) marginal sq.mm), 11-54% solitary, remainder in ra­ cells (heterogeneous III & 11), but in four dial and oblique multiples of 2-5(-7), occa­ shrub samples, multiseriate rays usually com­ sionally in clusters in C. campanulata (South posed of square to procumbent body ray cells China Agricultural University 2333) and C. and one to several rows of upright cells, the conradinae (Yunnan Academy of Forestry proportion of square to upright cells higher. 205), oval to round or angular, tangential Inflated secretory (?) cells (possibly contain­ diameter 23-65 (13-110) 11m, radial diam­ ing oil or mucilage) associated with ray pa­ eter 28-82 (15-120) 11m, walls I-311m renchyma were noted in Cerasus campanulata thick. Vessel element length 270-500 (130­ (Northeast Forestry University s.n.). 680) 11m. L/D ratio 6-13. Perforations ex­ Crystals present in all species except in C. clusively simple in oblique to almost horizon­ conradinae and C. tomentosa, sparse to abun­ tal end walls. Intervessel pits nonvestured, dant , prismatic and/or druses, medium-sized alternate, predominantly polygonal, rarely (druses) rarely to large (prismatic crystals) , in round to oval, 4-11 11m in diameter, with chambered axial parenchyma and/or ray slit-like apertures, apertures occasionally cells, one crystal in each more or less enlarg­ coalescent in C. clarofolia (HEFw 126), C. ed and sclerified chamber or ray cell ; chains pseudocerasus (FRIGw 463),C. serrulata of up to 14 chambers. Pith flecks mostly ab­ (HEFw 3160), C. serrulata var. sachalinen­ sent, but rare to common in e. campanulata sis and C. setulosa(HEFw 133). Vessel-ray (HEFw 5040), C. maximowiczii, C. serrula­ and vessel-parenchyma pits clearly smaller ta (CAFw 4865) and C. setulosa (CAFw 564). than intervessel pits and half-bordered with Traumatic gum ducts usually absent, but slightly reduced borders, 2-5 11m in diam­ present in C. cerasoides (FRIGw 1208) and eter. Helical thickenings weIl developed, usu­ C. yedoensis, large , in long tangential bands . ally coarse and widely spaced, throughout Notes: 1. The wood anatomy of Cerasus body of vessel elements. Gummy contents prostrata studied by Fahn et al. (1986) is present in some vessels of most species. rather different from the Chinese species Fibres 560-1400 (350-1620) 11m long, studied here. Cerasus prostrata has ring­ F/V ratio 1.9-3.4, medium thick-walled, porous wood, vessels mostly solitary, fibres with (minutely to) distinctly bordered pits of with distinctly bordered pits in radial and 2-5 11m in diameter, mainly confined to ra­ tangential walls , rays composed of square, dial walls, but simple to minutely bordered weakly procumbent and upright ceIls, and pits also present in thinner earlywood fibres crystals absent, which differs from the 14 in C. maximowicziiand C. yedoensis. Heli­ Chinese species studied. In addition, there cal thickenings mostly absent, but fine helical are also significant differences in some quan­ thickenings present in some latewood fibres titative features like vessel element size, fibre of C. pseudocerasus (Guizhou Forestry In­ length, and ray frequency and size, Schwein­ stitute s.n .) and some fibres of C. tomentosa. gruber (1990) also reported diffuse-porous to Parenchyma usually scanty, occasionally semi-ring-porous wood for Prunusprostrata fairly common, apotracheally diffuse and (or Cerasus prostrata). scanty paratracheal, in 3-5 (2-8)-celled 2. Ring-porous wood in Prunus cerasus strands. (or Cerasusvulgaris) was reported by Kribs Rays 5-14/mm, of two sizes except in (1968). However, diffuse-porous wood in two of four branch samples (C, maximo­ the species was reported by Greguss (1959) wiczii and C. tomentosa) where rays are 1-3­ and Oprea (1972). seriate: narrow and low rays 1(-2)-seriate, 3. Wood anatomically Cerasusis close to and wide and tall rays (3-)4-6(-7)-seriate. Prunuss.str. Both taxa share almost all qual­ Uniseriate rays 4-11 (1-30) cells high, com­ itative features. However, they can usually be posed of square and upright cells, or includ- distinguished from each other wood anatomi-

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cally: crystals are absent in Prunuss.str., and with slit-like apertures, apertures rarely co­ almost always present in Cerasus. alescent in one specimen of L. spinulosa 4. Librifonn fibres were reported by (FRIGw 1138). Vessel-ray and vessel-pa­ Schmidt (1941) in Prunus avium and by renchyma pits clearly smaller than intervessel Schweingruber (1990) in P. avium and P. pits and half-bordered, 2-4 J.1m in diameter. cerasus. Helical thickenings weil developed, usually 5. Cerasus clarofolia (368) is the only coarse and widely spaced, throughout body specimen in Prunus s.l. which has multi­ of vessel elements. Gummy contents present seriate rays of over 1.0 mm high. in some specimens of L. spinulosa. Fibres 910-1330 (500-1680) J.1m long, Laurocerasus Tourn. ex Duh. (Figs. 3, 4,11, F/V ratio (1.6-)1.9-2.9(-3.9), medium 12,20,32,47; Table 19) thick- to very thick-walled, with (minutely to) Evergreen trees or shrubs, very rarely de­ distinctly bordered pits of (2-)3-4 J.1m in ciduous, mainly from southern regions of the diameter, mainly confined to radial walls. Yellow River valley, especially South and Helical thickenings absent Southwest China. Parenchyma usually scanty, apotracheally Within Laurocerasus there are substantial diffuse and scanty paratracheal, in 3-4 (2-7)­ differences in wood structure. The following celled strands. two groups are recognised and described sep­ Rays 8-11 (6-14)/mm, 1-3(-4)-seriate arately. (but 4-seriate rays common in some samples, see Table 19). Uniseriate rays 3-6 (1-15) Group A cells high, composed of square and upright Material studied: L. phaeosticta (Hance) ceIls; multiseriate rays 290-570 (115-980) Schneid.: Guangxi, CAFw 16995; Guang­ J.UD high, composed of procumbent body cells dong, FRIGw 460; Guizhou, Guizhou For­ and 1-4 rows of square to upright marginal estry Institute s.n. - L. spinulosa (Sieb. & cells (heterogeneous III & 11). Zucc.) Schneid.: Guangdong, FRIGw 1138; Crystals, pith flecks and traumatic gum Taiwan, CAFw 7621; Guangxi, CAFw ducts absent. 15847; Fujian, CAFw 12662; Conghua, Gu­ angdong, South China AgriculturalUniversity Group B 1348; Guangxi, Guangxi Forestry College Material studied: Lifordiana (Dunn) Yu & s.n .; Jiangxi, CAFw 19173; Japan, CAFw Lu: Hainan, China, South China Agricultural 4866; Shaping, Sichuan, CAFw 367; Hunan, University 1285. - L. hypotriaha (Rehd.) Yu CAFw 14671. & Li: Guangxi, CAFw 15780. - L. undulata Growth rings usually distinct, marked by (D.Don) Roem.: Yunnan, CAFw 3604; Jiang­ rows of radially flattened latewood fibres. xi, CAFw 18043. - L. zippeliana (Miq.) Yu Vessels diffuse, in a weakly to distinctly ob­ & Li: Guangdong, CAFw 16680; Guang­ lique pattern in some samples, 73-199/sq. dong, CAFw 6572; Fujian, CAFw 9705. mm, 16-41% solitary, remainder mainly in Growth rings distinct to faint in L. fordi­ radial and oblique multiples of 2-6(-10), ana, L. hypotricha and L. zippeliana or faint multiples of over 4 common in two speci­ to indistinct in L. undulaia; mainly marked by mens of L. spinulosa (CAFw 19173 and 1-2(-4) rows of marginal zonate paren­ CAFw 15847), clusters occasionally present chyma bands, occasionally by weakly radial­ in some sampies. Vessels oval to round or Iy flattened latewood fibres as weil. Vessels weakly angular, tangential diameter 36-60 diffuse, in a radialpattern in most sampies (but (20-87) J.UD, radial diameter 53-79 (23­ in an oblique pattern in L.jordiana), 27-62/ 111) J.1m, walls 1-2 J.1m thick. Vessel ele­ sq.mm, 5-24% solitary, remainder mainly ment length 230-670 (130-810) J.1m. L/D in radial and oblique multiples of2-8, multi­ ratio 5-12(-17). Perforations exclusively ples of over 4 common in L. zippeliana, clus­ simple in oblique end walls. Intervessel pits ters occasionally present. Vessels oval to nonvestured, alternate, mostly polygonal, round, tangential diameter 61-79 (27-105) rarely round to oval, 5-10 J.1m in diameter, J.1m, radial diameter 78-108 (30-137) J.1m,

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Table 19. Variation in selected wood anatomical features of Laurocerasus.

234 5 6 7 8 9 10 11 12 13 14 GroupA L. phaeosticta CAFw 16995 - 116 21 36 410 5-7 1030 2.5 3-4 4 0.3 FRIGw460 o 73 21 60 410 5-8 1210 2.9 4-5 3 0.4 Guizhou s.n . o 154 26 42 410 6-9 980 2.4 4-5 4 0.3 L. spinulosa FRIGw 1138 95 24 49 570 5-9 1130 2.0 3- 5 3 0.6 CAFw 7621 - 111 41 50 500 6-10 1150 2.3 3-5 3 0.4 CAFw 15847 o 107 36 48 230 5-8 910 3.9 2-4 4 0.5 CAFw 12662 o 130 25 42 450 5-6 1020 2.3 4-5 2(-3) 0.3 1348 - 199 23 47 470 5-7 1330 2.8 4-5 3(-4) 0.5 Guangxi s.n . o 136 35 42 430 5-7 1240 2.9 3-4 3(-4) 0.3 CAFw 19173 o 125 21 40 670 6-10 1070 1.6 3-4 3(-4) 0.3 CAFw 4866 - 165 16 46 510 6-10 1020 2.0 4-5 4 0.5 CAFw 367 - 155 40 55 520 6-8 1010 1.9 3-5 3(-4) 0.5

Group B Li fordiana o 62 15 62 450 2-5 1220 2.7 2-5 3(-4) 0.5 D-A - + L. hypotricha R 53 24 75 360 2-4 1220 3.4 3-5 4-5 0.8 + + L. undulata CAFw 18043 27 24 60 620 2-5 1520 2.5 1-3 4 0.8 P-A + CAFw 3604 R 37 16 68 400 2-3 1080 2.7 3-5 10-12 0.9 P-AR + + L. zippeliana CAFw 16680 R 41 5 79 490 3-4 1400 2.9 3-4 5(-6) 0.7 + CAFw9705 R 47 24 61 460 2-4 810 1.8 4-6 4 0.7 +

I: Vessels in a weakly to distinctly oblique (0), or radial (R) pattern, orrandomly distributed (-); 2: Vessel frequency Usq.rnm); 3: Percentage of solitary vessels; 4: Average tangential vessel diameter (um); 5: Average vessel element length (11m); 6: Size of intervessel pits (um); 7: Aver­ age fibre length (um); 8: FIV ratio; 9: Size of pits in fibres (um); 10: Width of the widest rays (number ofcells); 11: Average multiseriate ray height (rnm); 12:Prismatic crystals (P), or druses (D) present in axial parenchyma cells (A), ray cells (R), in both (AR) or absent (-); 13: Traumatic gum ducts present (+) or absent (-); 14: Parenchyma bands present (+) or absent (-).

walls 2-4 11m thick. Vessel element length only in narrow vessel elements in one sampie 360-620 (270-880) 11m. LID ratio 5-7 (CAFw 3604) or absent in another one. Gum­ (-10). Perforations exclusively simple in ob­ my contents usua11y present lique end walls. Intervessel pits nonvestured, Fibres 810-1520 (650-1750) 11m, F/V alternate, mainly polygonal, 2-4(-5) 11m in ratio (1.8-)2.7-2.9, very thick-walled, with diameter, with slit-like apertures, apertures (minutely) to distinctly bordered pits of (2-) rarely coalescent in Lifordiana. Vessel-ray 3-5 11m in diameter, but simple to minutely and vessel-parenchyma pits similar to inter­ bordered pits also present in L. undulata vessel pits but half-bordered. Helical thick­ (CAFw 18043), mainly confined to radial enings weil developed, usually coarse and walls. Helical thickenings absent Gurnmy closely spaced, throughout vessel elernents, contents present in lumina of most fibres in but in L. undulata fine thickenings present L. zippeliana (CAFw 16680).

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Parenchyma moderately abundant, apotra­ the basal glands on leaves: section Phaeostic­ cheally diffuse, diffuse-in-aggregates, scanty tae including L. phaeostictaand L.fordiana; paratracheal and in 1-3-seriate marginal or seetion Laurocerasus including L. hypotricha, irregular zonate bands, in 3-5 (2-1O)-celled L. spinulosa, L. undulata, L. zippeliana, etc. strands. These two sections do not correspond with Rays 10-12 (9-13)/mm, mostly of two the two groups based on wood anatomy. sizes: narrow and low 1(-2)-seriate, and wide 5. Group A of Laurocerasus is the only and tall rays 3-5(-12)-seriate. Uniseriate one in Prunus s.l. where some species have rays 5-6 (1-15) cells high, composed of rays narrower than 4-seriate and of intergrad­ square and upright cells; multiseriate rays ing size only. 500-940 (150-1600) 11m high, composed of procumbent body cells and 1-5 rows of Padus Mill. (Figs. 31, 36; Table 20) upright to square marginal cells (heterogene ­ Material studied: P. brachypoda (Batal.) ous III & 11). Schneid.: Yinian, Yunnan, Yunnan Academy Crystals sparse in L.fordiana to abundant of Forestry 113, alt. 2000 m. - P. buergeri­ in L. undulata, prismatic (L. undulatai or ana (Miq.) Yu & Ku: Zhongtiaoshan, Shanxi, druses (L.fordiana), medium-sized to large, HEFw 135; Huangshan, Anhui, HEFw 3125, in chambered axial parenchyma cells in L. alt. 550 m. - P. grayana (Maxirn.) Schneid. : fordiana or both in ray and chambered (CAFw Muchuan, Sichuan, Sichuan For. Institute 3604) or non-ehambered (CAFw 18043) s.n.; locality unknown, CAFw 4104; Guang­ axial parenchyma cells in L. undulata, one dong, CAFw 17647. - P. maackii (Rupr.) crystal in each more or less enlarged chamber Kom .: Changbaishan, Jining, Qian Hong in L ifordiana and L. undulata (CAFw'3604) s.n. , diam. 18 mm; Northeast China, CAFw or 1-10 prismatic crystals in one non-enlarged 17308. -Po obtusata (Koehne) Yu & Ku: ray cell or axial parenchyma cell in L. undu­ Southeast China, HEFw 1182; Huangshan, lata (CAFw 18043); chains ofup to 17 cham­ Anhui, HEFw 1434, alt. 650 m; Sichuan, bers, crystals not observed in L. hypotricha HEFw 814; Sichuan, CAFw 812; Sichuan, and L. zippeliana. Pith flecks observed only CAFw 43. -P.perulata (Koehne) Yu & Ku: in L. undulata (CAFw 3604). Traumatic gum Sichuan, CAFw 1028. - P. racemosa (Lam.) ducts noted in L. hypotrichaand L. undulata Gilib.: Harbin, Heilongjiang, CAFw 17309; (CAFw 3604), in tangential bands. Northeast China, Northeast Forestry Univer­ Notes: 1. Metcalfe and Chalk (1950) stated sity s.n. - P. ssiori Schneid.: location un­ that Laurocerasus tends to have very definitely known, CAFw 4200. - P. wilsonii Schneid. : heterogeneous rays, frequently with 4 or more Sichuan, HEFw 787. rows ofupright marginal cells. In both groups Deciduous trees or shrubs, distributed of the genus in China, however, multiseriate throughout China, mainly from the Yangtze rays have 1-4(-5) rows of upright to square River valley, Shanxi and southern Gansu. marginal cells. Growth rings distinct, marked by rows of 2. Schweingruber (1990) noted that two radially flattened latewood fibres, in some species of Prunus s.l. in the sense of Ben­ samples by differences in vessel diameter be­ tham & Hooker (or Laurocerasus), Pi lauro­ tween latewood and subsequent earlywood as cerasus and P.lusitanica are characterised by well, Wood diffuse-porous, sometimes vessels in a distinctly radial pattern, as in ranging to semi-ring-porous. Vessels in a most Chinese species of Group B. weakly oblique pattern in P. brachypoda,P. 3. The two groups ofLaurocerasus differ grayana (CAFw 17647), P. maackii (Qian from each other mainly in growth ring boun­ Hong s.n.) and P. obtusata (HEFw 1434, daries , vessel pattern, vessel frequency, per­ CAFw 43 and HEFw 814), 40-246/sq.mm centage of solitary vessels, vessel diameter, (all below 170/sq.mm except in P. maackii, intervessel pits, vessel wall thickenings, axial Qian Hong s.n.), 5-54% solitary, remain­ parenchyma and ray size. der in radial, oblique and tangential multiples 4. Chinese Laurocerasusspecies were di­ of 2-7(-8), multiples of over 4 vessels fre­ vided by Yu (1986) into two sections using quent in P. perulata. Vessels oval to round or

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Tab1e 20. Variationin se1ected wood anatomicalfeaturesof Padus.

2 3 4 5 6 7 8 9 10 11 12

P. brachypoda R 90 24 48 380 7-9 1070 2.8 7(-8) 0.5 D-A P. buergeriana HEFw 135 - 170 54 38 380 5-8 790 2.1 6(-8) 0.5 D-R HEFw 3125 95 35 50 390 5-10 1060 2.7 5(-6) 0.5 P. grayana Sichuan s.n . 75 37 60 570 6-9 1220 2.2 6(-8) 0.5 D-A CAFw4101 - 143 44 49 370 5-7 960 2.6 7(-8) 0.8 + CAFw 17647 R 143 21 65 450 5-8 1070 2.4 8(-12) 0.5 + P. maackii Qian Hong s.n. R 246 21 30 300 4-7 740 2.2 2(-3) 0.5 CAFw 17308 - 112 40 53 550 5-8 970 1.8 2 0.4 Pi obtusata HEFw 1182 63 48 69 490 7-12 1270 2.6 4(-5) 0.6 HEFw 1434 R 110 32 50 490 5-11 1100 2.3 7 0.4 HEFw 814 R 40 47 63 440 6-12 1380 3.2 7(-8) 0.3 PD-AR ­ CAFw 812 52 47 59 460 7-10 1280 2.8 5(-6) 0.3 PD-AR ­ CAFw43 R 99 13 45 380 8-12 1220 3.2 7(-8) 0.4 PD-AR + P. perulata 79 10 55 390 7-10 1060 2.7 8(-10) 1.0 D-AR + P. racemosa CAFw 17309 - 140 32 45 260 5-8 790 3.0 4 0.2 Northeast s.n. - 139 36 42 310 4-5 840 2.7 4(-5) 0.2 P. ssiori 93 44 48 530 5-8 1030 1.9 5 0.4 P. wilsonii 48 5 75 520 8-10 1280 2.5 6(-7) 0.9

1: Vesse1s in a weakly to distictly radial pattern (R) or not (-) ; 2: Vesse1 frequency (/sq.mm); 3: Percentage of solitary vesse1s; 4: Average tangential vesse1 diameter (J.lII1); 5: Average vesse1 element 1ength (J.lII1); 6: Size ofintervesse1 pits (J.lII1); 7: Average fibre length (J.lII1); 8: F/V ratio; 9: Width of the widest rays (number of cells); 10: Average mu1tiseriate ray height (mm); 11: Prismatic crystals (P), druses (D) or both (PD) present in axial parenchyma ce1s (A), ray cells (R), in both (AR) or absent (-); 12: Traumatic gum duets present (+) or absent (-). angular, tangential diameter 30-75 (l7-100) ed with reduced borders, 2-5 um in diam­ um, radial diameter 39-85 (20- 160) J.lII1, eter. Helical thickenings weil developed, usu­ walls 1-3 um thick. Vessel element 1ength ally coarse and wide1y spaced, throughout 260-570 (140-730) um, LID ratio 5-11. body of vessel elements. Gummy contents Perforations exclusively simple in oblique to present in most species. almost horizontal end walls. Intervesse1 pits Fibres 740-1380 (390-1770) um long, nonvestured, alternate, predominantly poly­ FIV ratio 1.8-3.0, very thin-to very thick­ gonal, occasionally round to oval, 4-12 um walled, mostly with minutely to distinctly in diameter, with slit-like apertures, apenures bordered pits of 2-51lm in diameter, mainly rarely coalescent in P. brachypoda, P. buer­ confined to radial walls, minutely bordered geriana (HEFw 135), P. grayana (Sichuan pits present in thinner earlywood fibres of Forestry Institute s.n.), P. maackii (Qian some species. Helical thickeningsabsent. Hong s.n.), P. perulata and P. ssiori. Ves­ Parenchyma usually scanty, apotracheally sel-ray and vessel-parenchyma pits clearly diffuse and scantyparatracheal,rarelydiffuse­ smal1er than intervessel pits and half-border- in-aggregates, in 3-4 (2-8)-celled strands.

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Rays 4-14 (3-15)/mm, of two sizes Notes: 1. Metealfe and Chalk (1950) men­ (except in two branch samples ofP. maackii tioned the occurrence ofring-porous or semi­ with 1-2(-3)-seriate rays): narrow and low ring-porous woods in some species of the rays 1(-2)-seriate, and wide and tall rays (2-) genus . But all the samples we examined are 4-8(-12)-seriate. Uniseriate rays 3-8 (1-22) diffuse-porous to semi-ring-porous. Diffuse­ cells high, composed ofprocumbent cells on­ porous woods in Padus species were also ly, or with one row of square to upright mar­ reported by Oprea (1972). ginal cells; multiseriate rays 230-970 (70­ 2. Padus stands out in Prunus s.l. (in the 2900) um high, composed of procumbent the sense of Kalkman) on account of its cells only, or sometimes with one row of mainly homogeneous rays. square marginal cells (mainly homogeneous, 3. Foraminate or reticulate perforations in sometimes heterogeneous III also present), some species of the genus were recorded by Crystals sparse to common in some spe­ Metcalfe and Chalk (1950), However, multi­ cies, druses and prismatic crystals, usually ple perforations were not observed in our medium-sized, in chambered axial parenchy­ material. ma, ray cells or both, one crystal in each more or less enlarged chamber or ray cell, Prunus s.str. L. (Table 21) chains of up to 14 chambers, crystals not Material studied: P. domestica L.: location observed in P. buergeriana (HEFw 3125), P. unknown, FHOw 3079. - P. salicinaLindl.: grayana(CAFw 4101 and CAFw 17647), P. Zhongtiaoshan, Shanxi, HEFw 132; Huo­ maackii, P. obtusata (HEFw 1182 and HEFw shan, Anhui, He Yunhe s.n., diam, 20 mm, 1434), P. racemosa, P. ssioriand P. wilsonii. alt. 200 m; Huangshan, Anhui , HEFw 3121, Pith flecks rare to common in P. brachypoda, alt. 400 m; Guizhou, Guizhou Forest Insti­ P. grayana(CAFw 17647 and CAFw 4101), tute s.n.; Guangxi, CAFw 15874; Guang­ P. maackii (Qian Hong s.n.) and P. obtusata dong , FRIGw 1241. - P. spinosa L.: loca­ (CAFw 43). Traumatic gum ducts rare to tion unknown, FHOw 2822 . - P. ussuriensis common in P. grayana (CAFw 4101 and Kov. & Kost. : Jining, CAFw 18077. CAFw 17647) , P. obtusata (CAFw 43) and Deciduous small trees or shrubs distributed P. perulata, small to large, in long or short throughout China, widely cultivated as impor­ tangential bands or diffuse. tant fruit trees and ornamentals.

Table 21. Variation in selected wood anatomical features of PrunusS.str.

1 2 3 4 5 6 7 8 9 P. domestica 150 45 38 4-8 5(-6) 0.5 P. salicina HEFw 132 180 20 47 330 6-10 730 2.2 4(-5) 0.5 He Yunhe s.n . 179 53 30 300 5-8 580 1.9 6(-7) 0.3 HEFw 3121 154 47 36 270 5-8 800 3.0 4(-5) 0.4 Guizhou s.n. 163 39 39 280 4-6 870 3.1 8(-9) 0.6 CAFw 15874 55 67 35 550 5-8 1280 2.3 8(-9) 0.5 FRIGw 1241 110 56 38 310 5-10 740 2.4 6(-8) 0.5 P. spinosa 115 49 38 5-9 4(-5) 0.7 P. ussuriensis 280 48 34 560 9-11 1200 2.1 2 0.3 1: Vessel frequency (/sq.mm); 2: Percentage of solitary vessels ; 3: Average tangential vessel diameter (um); 4: Average vessel element length (um); 5: Size of intervessel pits (um); 6: Aver- age fibre length (um): 7: F/V ratio ; 8: Width of the widest rays (number of cells); 9: Average multiseriate ray height (um).

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Growth rings distinct, marked by rows of Pygeum Gaertn. (Figs. 2, 5, 33, 44) radially flattened 1atewood fibres, in a few Material studied: P. topengii Merr.: Guang­ species by differences in vessel diameter be­ dong, FRIGw 22; Hainan, CAFw 14370; tween latewood and subsequent ear1ywood Guangdong, CAFw 6571. as well. Wood usual1y diffuse-porous (but Evergreen trees or shrubs, mainly from semi-ring-porous in one specimen of P. sali­ South to Southwest China. cina). Vesse1s 1l0-280/sq.mm, 20-56 Growth rings usually faint, marked by (-67)% solitary, remainder in radial, oblique zonate parenchyma bands. Vessels diffuse, and tangential multiples of 2-6(-8), oval to 8-11/sq.mm, 20-55% solitary, remainder round or weakly angular, tangential diameter mainly in radial multiples of 2-5, oval to 30-47 (20-60) um, radial diameter 35-62 round, tangential diameter 100-123 (46-170) (22-80) um, walls 1-211m thick. Vessel ele­ um, radial diameter 141-160 (60-218) um, ment length 270-560 (180-760) um, L/D walls 2-4 um thick. Vessel element length ratio 6-10(-16). Perforations exclusively 530 (160-790) um. L/D ratio 4-5. Perfora­ simple in oblique end walls. Intervessel pits tions exclusively simple in oblique to hori­ nonvestured, alternate, polygonal or round to zontal end walls. Intervessel pits nonvestured, oval, 4-11lJ.ffiin diameter, with slit-like aper­ alternate, mainly polygonal, rarely round to tures, apertures occasionally coalescent in P. oval, 2-5 um in diameter, with slit-like to domestica and P. salicina (CAFw 1241). Ves­ coalescent apertures. Vessel-ray and vessel­ sel-ray and vessel-parenchyma pits clearly parenchyma pits similar to intervessel pits but smaller than intervessel pits and half-border­ half-bordered, occasionally unilaterally com­ ed with reduced borders, 2-5 um in diam­ pound or elongate. Helical thickenings fine eter. Helical thickenings well developed and and closely spaced, confined to narrow ves­ widely spaced, throughout body of vessel sel elements or to tails. Gummy contents ab­ elements. Gummy contents usually present. sent. Fibres 580-1280 (290-1550) um long, Fibres 1060-1310 (720-2010) um long, F/V ratio 1.9-3.1, medium thick- to very F/V ratio 2.0-2.5, very thin-to medium­ thick-walled, with (minutely to) distinctly thick-walled, usually with minutely to dis­ bordered pits, (2-)3-5 um in diameter, tinctly bordered pits of 2-4 um in diameter, mostly confined to radial walls. Helical thick­ confined to radial walls. Helical thickenings enings absent. absent. Parenchyma scanty, apotracheally diffuse Parenchyma moderately abundant, apotra­ and scanty paratracheal, in 3-4 (2-6)-celled cheal1y diffuse and scanty paratracheal to strands . vasicentric, also in narrow irregular zonate Rays 7-13 (6-16)/mm, of two sizes: nar­ (sometimes marginal) bands of up to 10 cells row and low rays 1(-2)-seriate, and wide wide, in 4-6 (2-11)-celled strands. and tall rays 3-8(-9)-seriate. Uniseriate rays Rays 4-8 (2-9)/mm, 1-4(-5)-seriate, 3-7 (1-20) cells high, composed of square more or less of two sizes: uniseriate rays, and to weakly procumbent cells and one to sev­ (2-)3-4(-5)-seriate. Uniseriate rays 5-8 eral rows of upright cells; multiseriate rays (1-13) cells high, composed of square to up­ 306-710 (130-1440) um high, composed right cells; multiseriate rays 380-520 (170­ of procumbent body cells and 1-5 rows of 850) um high, composed of procumbent square (to upright) marginal cells (heterogen­ body ray cells and 1-4 rows of square to up­ eous III & 11), but in one branch sample of right marginal cells (heterogeneous III and Il), P. salicina (He Yunhe s.n.), multiseriate rays Crystals and pith flecks absent. Traumatic composed of square to procumbent body ray gum ducts present in one sample (FRIGw cells and rows of upright marginal cells. Pro­ 22), in long tangential bands. portion of square and upright ray cells higher Notes : 1. Vasicentric tracheids , fonning in juvenile wood than in mature wood. narrow sheaths around the vessels, were noted Crystals absent. Pith flecks common in P. by Desch (1954) in Pygeum p.p. No vasicen­ salicina (HEFw 3121 and Guizhou Forest tric tracheids, however, were observed in the Institute s.n.). Traumatic gum ducts absent. Chinese specimens examined.

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2. Rao & Purkayastha (1972) stated that families, as indicated in the survey. However, Prunus s.l. (in the sense of Bentham & the Spiraeoideae and Rosoideae are similar to Hooker) and Pygeum are wood anatomically each other in wood strucrure and the two sub­ quite distinct and that Pygeum can be distin­ families cannot be distinguished. The main guished from Prunus s.l. by the presence of wood anatomical differences between the vasicentric parenchyma and absence of heli­ Spiraeoideae & Rosoideae, Maloideae and cal thickenings. However, helical thickenings Prunoideae are summarised in Table 22. The are present in narrow vessel elements or tails most appreciable differences among the fea­ of elements in Pygeum topengii. Absence of tures listed are in quantitative and qualitative helical thickenings in Pygeum henryi was re­ ray characters. Other discriminating features ported by Luo (1989). are percentage of solitary vessels, vessel wall 3. The present study indicates that Pygeum thickenings, and attributes of ground tissue differs from all the taxa in Prunuss.l. (except fibres, crystals and axial parenchyma. Group B of Laurocerasus) in both qualitative Within most subfarnilies wood anatomi­ and quantitative features. The genus is char­ cal features are relatively homogeneous. The acterised by zonate parenchyma bands, a low Prunoideae, however, are rather diverse. vessel frequency and a high degree of vessel The Spiraeoideae & Rosoideae show a grouping, wide vessel elements, intervessel limited diversity in wood anatomical features, pits similar to vessel-ray and vessel-paren­ and the seven genera studied can be separated chyma pit, and abundant parenchyma. wood anatomically. Spiraea is unique for its two types of fibres; Rubus is exceptional for Notes on Prunus s.l. its higher degree of vessel grouping and in­ 1. Metcalfe and Chalk (1950) reported that distinct growth ring boundaries; Rosa is noted rays in Prunus s.l. (in the sense of Bentham for its ring-porous wood as weil as irregular & Hooker) are more homogeneous than in vessel wall thickenings; Kerria is character­ Laurocerasus, that there are rarely more than ised by scalariform to opposite intervessel 1-3 marginal rows in Prunus s.l., and that pits; Stephanandra can be separated because it marginal ray cells are square rather than up­ lacks the features of the genera mentioned right in Laurocerasus. These differences do above. All the five genera mentioned are not hold true in the Chinese species studied. characterised by rays of two distinct sizes: 2. Traumatic gum ducts in Prunus s.l. (in wide and tall rays (3-)4-6(-16)-seriate, 1.0 the sense of Bentham & Hooker) were also to 25.0 mm high, composed of square or recorded by Barajas Morales (1980), Metcalfe square to procumbent body cells and upright & Chalk (1950), Record & Hess (1943) and marginal cells, usually also with weakly dif­ Tang (1976). In addition, both Suzuki (1984) ferentiated sheath cells. In addition, they also and Cevallos-Ferriz & Stockey (1990) recog­ share some features including more or less nised the incidence of traumatic gum ducts in wavy growth ring boundaries, fibre pits usu­ fossil woods of Prunus s.l . (in the sense of ally densely spaced in radial walls, inter­ Bentham & Hooker). vessel pits similar to vessel-ray pits, scanty axial parenchyma and medium-sized, non­ Wood anatomy and classification of chambered prismatic crystals sometimes in the Rosaceae from China non-enlarged ray cells. Exochorda differs from the genera mentioned above in most The classification of the Rosaceae has wood anatomical features, as shown in Table been a matter of continued debate. Based on 23, and resembles the Prunoideae. Sorbariais this wood anatomical study of Chinese Rosa­ to a lesser extent different from the other gen­ ceae, only abrief and tentative discussion era of the Spiraeoideae & Rosoideae , mainly will be given. Wider taxonomie implications in ray characters (composition and height) will be reviewed in a subsequent paper based and LID ratio (Table 23). on a worldwide wood anatomical survey. The Maloideae constitute a quite homo­ There are appreciable differences in some geneous wood anatomical group. The fifteen wood anatomical features between the sub- genera studied are very similar to each other.

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Table 22. Main differences in wood anatomical features between the Spiraeoideae & Rosoideae, Maloideae and Prunoideae from China.l

Spiraeoideae Maloideae Prunoideae & Rosoideae

Vessels ~ 60% solitary + (-) + - (+) L/D ratio (common range) 8-18 10-26 4-14 Intervessel pits polygonal + (-) -(+) + Vessel-ray pits clearly smaller than intervesselpits - (+) + (-) Helical wall thickenings -(+) + (-) + (-) Gummy contents - (+) +/- + (-) Fibre pits densely placed in radial walls +/- Fibre pits mainly confined to radial walls -(+) + (-) F/V ratio (common range) 1.4-2.6 1.4-2.6 1.8-3.4 Parenchyma abundant -(+) + - (+) Marginalparenchyma -/+ The widest rays ~ 4-seriate + -(+) + (-) Ray height (mm) (0.4-)1.0-25.0 s 0.4 0.3-1.0 Body ray cells procumbent -/+ + + Crystals in ray cells +/- +/- Crystals in enlarged and/or chambered parenchyma + (-) +/- Druses +/- Traumatic gum duets -/+

Table 23. Main differencesin wood anatomicalfeatures between Exochorda, Sorbaria and remaining genera in the Spiraeoideae & Rosoideae from China.l

remaining Exochorda Sorbaria Spireoideae& Rosoideae

Growth ring boundaries wavy +/- + (-) Ring-porosity + -(+) L/D ratio (only for diffuse-porous species) 6-8 (6-)8-19 Helical wall thickenings + -(+) Fibre pits densely placed in radial walls +/- + (-) Fibre pits mainly confined to radial walls + Helical thickeningsin fibres + -(+) Parenchyma scanty + (-) Ray height s 1.0 mm + + Body ray cells procumbent + + -/+ Crystals + +/-

1) () = charaeter of sporadie occurrence.

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Table 24. Wood anatomical diversity in the Prunusalliance in China.l

~ .s<.J 0:: ~ ~ '() ~ ::l '"0:: ~ e '" C>() ...'" ~ ::l '1;3 ::l E' ~ ~ ... ;:... '

1) () = character of sporadic occurrence.

The subfamily is characterised by a high per­ most one row of square (to upright) marginal centage of solitary vessels, ground tissue cells, 2) Dichotomanthus, Cotoneaster, Stran­ composed exclusively of fibre-tracheids with vaesia,Photinia, Eriobotrya, Raphiolepisand distinctly bordered pits of 3-9 um in diam­ Chaenomeles, where multiseriate rays are eter, in radial and tangential walls, relatively composed ofprocumbent body cells and 1- 6 abundant axial parenchyma, 1-3(-5)-seriate rows of square to upright marginal cells. rays, not more than 004 mm high and almost Wood anatomically the Prunoideae are the always of intergrading sizes only and large least homogeneous in the farnily. Neverthe­ prismatic crystals in enlarged, sclerified and/ less, all genera in the subfarnily (except Prin­ or chambered axial parenchyma cells. Wood sepia) share a few wood anatomical features, anatomically it is impossible to key out most [e.g. less than 55(-60%) of the vessels soli­ individual genera. One appreciably variable tary, fibre pits mainly confined to the radial character within the subfarnily is ray compo­ walls, F/V ratio 1.6-4.0, axial parenchyma sition. In addition, some differences exist in mostly scanty, usually rays of two sizes, and vessel wall thickenings. Based on ray com­ multiseriate rays 0.3-1.0 mm high]. Prin­ position, the fifteen genera studied can be sepia is somewhat different from Prunus s.l. separated into two groups : 1) Sorbus, Pyrus, (in the sense of Kalkman 1965) in fibre pit Malus, Crataegus, Amelanchier, Cydonia, distribution, percentage of solitary vessels, and Docynia, where multiseriate rays are com­ and ray characters. The seven groups in Chi­ posed of procumbent cells only, or have at nese Prunus s.l. can be recognised based on

Downloaded from Brill.com10/09/2021 05:06:33PM via free access Zhang & Baas - Wood anatomy of Rosaceae from China 85 wood anatomical features, as shown in Table Laurocerasus differ from each other in 24. Amygdalus andArmeniaca are unique for growth ring boundaries, vessel pattern, ves­ their ring-porous-wood. These two genera seI frequency and degree of vessel grouping, are very similar in other features as weIl and vessel diameter, vessel-ray pits versus inter­ thus can be combined into one wood anatom­ vessel pits, vessel wall thickenings, axial pa­ ical group. Padus stands out on account of its renchyma and ray width. As shown in Table mainly homogeneous rays, while the rays in 24, Group A of Laurocerasus is characterised the remaining genera of Prunus s.l. are het­ by features which are common to most spe­ erogeneous with 1-5 rows of square to up­ eies of the traditional Prunus s.l. (in the right marginal cells. Prunuss.str. and Cera­ sense of Bentham & Hocker). On the other susshare most wood anatomical features, but hand, Group B is wood anatomically quite can, at least in China, usually be distinguished distinct from Prunus s.l. (in the sense of from each other: crystals are absent in Prunus Bentham & Hocker), but much closer to s.str., while they are almost always present Pygeum. However, there are still appreciable in Cerasus. Pygeum differs greatly in wood differences between Laurocerasus group B anatomy from all the taxa mentioned above and Pygeum (e.g. vessel diameter or LID (Table 24). The two groups recognised in ratio).

Key to tbe gmera of Rosaceae fI'om China

As shown in the following dichotomous key, all genera belonging to the Spiraeoideae, Rosoi­ deae and Prunoideae studied from China are distinguishable wood anatomically. However, most genera in the Maloideae cannot be differentiated. It should be kept in mind that the key should only be used for the Rosaceae from China.

1a. Multiseriate rays on average higher than 1.0 mm 2 b. Multiseriate rays on average lower than 1.0 mm ...... 7

2a. Ground tissue composed of two types of fibres, including libriform fibres with simple to minutely distinctly bordered pits of 1-2 um Spiraea b. Ground tissue composed exclusively of fibres with distinctly bordered pits of 3-5(-7) J.I.l11. ••••••••••••••••••••••••••••••••••••••••••••••••••••••• 3

3a. Wood ring-porous, vessel wall thickenings present Rosa b. Wood diffuse-porous or intergrading to semi-ring-porous,vessel wall thiekenings absent 4

4a. Multiseriate rays on average lower than 5.0 mm ...... 5 b. Multiseriate rays on average higher than 5.0 mm ...... 6

5a. Crystals present Stephanandra b. Crystals absent Rubus p. p. (R. corchorifolius, R. biflorus)

6a. Intervessel pits alternate Rubus p. p. b. Intervessel pits alternate and occasional1yalso scalariform to opposite Kerria

7a. Rays 1-3(-4)-seriate (but 4-seriate rays never common), multiseriate rays not higher than 0.4(-0.5)mm...... 8 b. The widest rays not narrower than 4-seriate, multiseriate rays 0.3-1.0 mm high . . . . 11

8a. Multiseriate rays composed of procumbent body cells and 1-6 rows of square to upright marginalcells ...... 9 b. Multiseriate rays composed ofprocumbent cells only, or sometimes with one row of square marginal cells Amelanchier, Crataegus , Cydonia, Docynia, Malus, Pyrus, Sorbus

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9a. Fibre pits main1y confined to radial walls Laurocerasus A p.p . b . Fibre pits common in radial and tangential walls ...... 10

lOa. Helical thickenings in fibre-tracheids absent Eriobotrya, Photinia A p.p. b. He1icalthickenings in fibre-tracheids present Chaenomeles, Cotoneaster, Dichotomanthus, Photinia A p.p., Prinsepia p.p. (P. sinensis), Pyracantha, Raphiolepis, Stranvaesia

lla. Marginal or zonate parenchyma bands present 20 b. Parenchyma bands absent ...... 12

12a. Over 65% of the vessels solitary (or latewood vessels solitary for ring-porous species) 13 b. Less than 60% of the vessels solitary (or latewood vessels solitary) ...... 16

13a. Crystals present in chambered and enlarged axial parenchyma cells .. .:.. .. Photinia B b. Crystals present in non-enlarged ray cells or absent ...... 14 14a. Vessel wall thickenings absent Sorbaria b. Vessel wall thickenings present ...... 15 ISa. Wood ring-porous to semi-ring-porous Exochorda b. Wood diffuse-porous to semi-ring-porous Prinsepia p.p . (P. utilis) 16a. Wood ring-porous Amygdalus, Armeniaca b. Wood mostly diffuse-porous , or rarely grading to semi-ring-porous 17 17a. Multiseriate rays composed of procumbent cells only, or sometimes with one row of square marginal cells ' Padus b. Multiseriate rays composed of procumbent body cells and 1-4(-6) rows of square to up- right marginal cells ...... 18 18a. Crystals present Cerasus b. Crystals absent 19 19a. Rays 1-4-seriate, vessels usually in a more or less oblique pattern . Laurocerasus A p.p. b. Widest rays wider than 4-seriate, vessels usually in random arrangement .. Prunuss.str. 20a. Tangential vessel diameter on average smaller than 80 lUD LaurocerasusB b. Tangential vessel diameter on average larger than 90 um Pygeum

Aclmow1edgements of Wood Industry, Yunnan Academy of The authors gratefully acknowledge the Forestry; Mr. Su Zhonghai, Department of Royal Dutch Academy of Seiences for the Forest Products, Guangdon Institute of For­ financial support, with which the first author estry ; Mr. Chen Jianchao, Department of could undertake this study at the Rijksher­ Forestry, South China Agricultural Univer­ barium/ Hortus Botanicus of Leiden Univer­ sity; Prof. Xie Fuhui, Guangxi Forestry sity. Thanks are due to all who generously College; Mr. Zhu Yiqian, Sichuan Institute of provided us with wood sampies or micro­ Forestry; Prof. Ho Tienxian, Department of scopic slides: Prof. Ke Bingfan, Prof. Wei Biology, Sun Yixian University (Guang­ Guangyang and Mrs . Jiang Zehui, Institute zhou) ; Prof. Dai Chengyue, Department of of Forest Products, Anhui Agricultural Col­ Fores t Indu stry, Northeast China Forestry lege; Mr. Liu Peng, Mr. Yang Jiaju and Mr. University; Curator of Guizhou Institute of Hong Diaoyian, Wood Anatomy Division, Forestry; and our sincere thanks are also due Institute of Wood Industry, Chinese Acad ­ to all who kindly collected necessary shrubby emy of Forestry; Mr. Luo Niangcai, Institute specimens in the field for this study: Dr. Qian

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Hou, Sheng Yang Institute of Applied Ecol­ Barajas Morales, J. 1980. Anatomia de ma­ ogy of Academia Sinica; Mrs . Liu Xiumei, deras de Mexico. No. 3. Diez espeeies del Department of Forestry, Anhui Agricultural bosque caducifolio de las cercanias de Xa­ College: Mr. Ye Shuyou, Huangshan Fores­ lapa, Ver. Mexico. Biotica 5: 23-40. try School; Mr. He Yunhe and Mr. Huang Bentham, G. & J.D. Hooker. 1865. Genera Chenlin, Department of Forestry, Anhui Agri­ Plantarum. London. cultural College. We are indebted to Prof. Brazier, J.D . & G.L. Franklin. 1961. Iden­ Dr. C. Kalkman (Leiden) for his review of tification of hardwoods. A microscope the manuscript, Ms. Bertie Joan van Heuven key. For. Prod. Res . BuH. No. 46, Lon­ (Leiden) for her skilful technical assistance don. and Prof. Dr. E.A. Wheeler (Raleigh) for Burgess, P.F. 1966. Timbers of Sabah. Sa­ useful discussion. bah For. Res . No. 6, Sabah. The first author wishes to record his spe­ Burgh.J. van der. 1978. Hölzer aus dem Pli ö• cial gratitude to Prof. Ke Bingfan (Hefei) for zän der Niederrheinischen Bucht . Fort­ his encouragernent, support and help through­ schr.Geol. Rheinl. u.Westf. 28: 213-275. out this study . Bykova, N.B. 1966. The anatomical struc­ ture of shoots in some species of Amyg­ Refurences dalus . Bot. Zhurnal 51: 517-525. (In Baas, P. 1973. The wood anatomieal range Russian.) in Ilex (Aquifoliaceae) and its ecological Carlquist, S. 1985. Vasicentric tracheids as a and phylogenetic significance. Blumea 21: drought survival mechanism in the woody 193 -258. flora of Southern California and similar Baas, P. 1985. A new multilingual glossary regions; review of vasicentric tracheids. of terms used in wood anatomy? IAWA Aliso 11: 37-68. Bull. n. s. 6: 83. Carlquist, S. 1986. Terminology of irnper­ Baas, P. 1986. Terminology of imperforate forate tracheary elements. IA WA BuH. tracheary elements : in defence of libriform n. s. 7: 75-81. fibres with minute bordered pits. IAWA Carlquist, S. 1988a. A new pathway in evo­ Bull. n. s. 7: 82-86. lution of irnperforate tracheary elements. Baas, P., P.M. Esser., M.E.T. van der Wes­ Aliso 12: 103-118. ten & M. Zandee. 1988. Wood anatomy Carlquist, S. 1988b. Comparative wood anat­ of Oleaceae. IAWA Bull. n. s. 9: 103­ omy. Springer Verlag, Berlin, Heidelberg, 182. New York, London, Paris, Tokyo. Baas, P., Lee Chenglee, Zhang Xinying, Cui Carlquist, S & D.A. Hoekman. 1985. Eco­ Keming & Deng Yuefen . 1984. Some ef­ logical wood anatomy of the woody fects of dwarf growth on wood structure. Southern Califomia flora. IA WA BuH. IAWA BuH. n.s. 5: 45-63. n.s. 6: 319-347. Baas, P. & F.H. Schweingruber. 1987. Bco­ Cevallos-Ferriz, S.R.S . & R.A. Stockey. logical trends in the wood anatomy of 1990. Vegetative remains of the Rosaceae trees, shrubs and climbers from Europe. from the Princeton chert (Middle Eocene) IAWA BuH. n.s, 8: 245-274. of British Columbia. IAWA Bull. n. s . Baas, P., P. Werker & A. Fahn . 1983. Some 11: 261-280. ecological trends in vessel characters. Challice, 1.S . 1974. Rosaceae chemotaxon­ IAWA BuH. n.s. 4: 141-159. omy and the origins of the Pomoideae. Baas,P. & Zhang Xinying. 1986. Wood Bot. J. Linn. Soc. 69: 239-259. anatomy of trees and shrubs from China: ChaHice, J.S. 1981. Chemotaxonomic stud­ I. Oleaceae. IA WA BuH. n.s, 7: 195-220. ies in the family Rosaceae and the evolu­ Bailey, LW. 1933 . The cambium and its tionary origins of the subfarnily Maloideae. derivative tissue No. VII. Structure, dis­ Preslia 53: 289-304. tribution, and diagnostic significance of Cheng, 1.Q. 1980. Chinese tropical and sub­ vestured pitsin Dicotyledons. 1. Amold tropical timbers. Science Press, Beijing. ArOOr. 14: 259-272. (In Chinese.)

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