IAWA Journal, Vol. 19 (1),1998: 43-97

SYSTEMATIC WOOD ANATOMY OF CORNACEAE AND ALLIES by

S. Noshiro l & P. Baas2

SUMMARY

The wood anatomy of Comaceae, Alangiaceae, Garryaceae, and Nyssa­ ceae constituting the Comales in the sense of Cronquist (1981, 1988) is described in great detail and subjected to a cladistic analysis. A micro­ scopic identification key to the woods studied is given. The alliance in­ cludes seventeen genera, mostly of trees and shrubs, very rarely herbs. Although wood anatomically fairly homogeneous, variation exists in both qualitative and quantitative characters. Some of the latter show distinct latitudinal trends within individual genera, and character states have only been recognised taking their latitudinal dependencies into ac­ count. The character states ultimately recognised in these continuously varying quantitative characters coincide with intergeneric or intersec­ tional gaps. The cladistic analysis based on a datamatrix with twenty­ one characters (Table 3) and using Cereidiphyllum, Daphniphyllum, and Hamamelis as outgroups yielded a strict consensus tree with a quadrichotomy with two monophyletic clades, Hydrangea panieulata (a representative of the closely allied Hydrangeaceae) and Daph­ niphyllum (Fig. 81). One weakly supported clade includes Alangium, Camptotheea, Cornus, Curtisia, Davidia, Diplopanax, Mastixia, and Nyssa without any robust lineages among them. The other genera, Ara­ lidium, Aueuba, Corokia, Garrya, Griselinia, Helwingia, Melanophylla and Toricellia, constitute a second, well-supported clade. Two Hydran­ gea taxa included in the analysis nest in the second clade and a basal branching respectively. The wood anatomical diversity pattern thus supports a family concept of Comaceae including Cornus, Curtisia, Diplopanax, Mastixia, Alangiaceae, and Nyssaceae, and exclusion of the genera in the other clade. There is remarkable agreement between some of these wood anatomical r~sults and recent cladistic analyses of rbeL sequences by Xiang and co-workers. The infrageneric classifica­ tion of Cornus, Alangium and Nyssa is also discussed. Key words: Comaceae, Alangiaceae, Garryaceae, Nyssaceae, Hydran­ gea, wood anatomy, latitudinal trends, cladistic analysis.

1) Forestry and Forestry Products Research Institute, Tsukuba Norin, P.O. Box 16, Ibaraki 305, Japan. 2) Rijksherbarium/Hortus Botanicus, P. O. Box 9514, 2300 RA Leiden, The Netherlands.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Table 1. Systematic treatment of the Comaceae alliance. 1:1:

Cronquist Harms Wangerin Me1chior Hutchinson Takhtajan Eyde Thorne 1982 1898 1910 1967 1967 1997 1988 1992 Cornales Cornaceae Cornaceae U mbelliflorae Araliales Comales Cornaceae Cornales Comaceae Aralidium (Araliaceae) Araliaceae (Aralidiales) Aralidiaceae Aucuba * * * * (Aucubales) Aucubaceae Cornus * * * * * * * Corokia * * * * (Hydrangeales) (Hydrangeales) Curtisia * * * * Curtisiaceae Curtisiaceae Griselinia * * * * (Griseliniales) (H ydrangeales) Helwingia * * * Araliaceae (Helwingales) (Araliales ) Kaliphora * * * (Hydrangeales) (Hydrangeales) Mastixia * * * * Mastixiaceae * * Melanophylla * * * * (Hydrangeales) (Hydrangeales) Toricellia * * * * (Toricelliales) (Araliales) (Diplopanax) Araliaceae Mastixiaceae *1) * Nyssaceae Nyssa * Nyssaceae Nyssaceae Nyssaceae Nyssaceae * * Camptotheca * Nyssaceae Nyssaceae Nyssaceae Nyssaceae * * Davidia * Nyssaceae Davidiaceae Nyssaceae Davidiaceae * * Garryaceae ......

Downloaded fromBrill.com10/10/2021 08:45:33PM Garrya (Garryales) * Garryaceae Garryaceae Garryaceae Garryaceae ~ Alangiaceae ;J> '-< Alangium * Alangiaceae Alangiaceae Alangiaceae Alangiaceae Alangiaceae 0 Araliaceae Araliaceae Vitaceae at= Umbelliferae Caprifoliaceae Gunneraceae f-. Haloragaceae Eucommiaceae ~ ...... Icacinaceae \D Cardiopteridaceae -----...... ------_ ... _-- '-' * = inclusion in Cornaceae, () = exclusion from Cornaceae or Cornales......

via freeaccess \D \D 1) Eyde & Xiang 1991. 00 Noshiro & Baas - Systematic wood anatomy of Comaceae 45

INTRODUCTION

Comaceae and allies constitute a group oftrees and shrubs (very rarely herbs) which has its main distribution in the northem hemisphere. The circumscription of this alli­ ance has been in constant dispute and varies greatly between systematists (Table 1; cf. also Xiang et al. 1993). Harms (1898) recognised 15 genera in seven subfamilies in his treatment of Comaceae. Wangerin (1910) divided Comaceae into four separate farnilies, Comaceae, Nyssaceae, Alangiaceae, and Garryaceae. In more recent years Cronquist (1981) adopted the family circumscription of Harms (1898) in his Comales and regarded the four families ofWangerin (1910) to constitute the order. Cronquist (1988) later included Nyssaceae in Comaceae following the opinions of Eyde (1988). Hutchinson (1967) included Comaceae, Alangiaceae, Garryaceae, and Nyssaceae to­ gether with Araliaceae and Caprifoliaceae in his Araliales, and Melchior (1964) in­ cluded the four families with Araliaceae and Umbelliferae in his series Umbelliflorae. Dahlgren (1980) and Thome (1992) have the broadest concept of the Comales. Dahlgren included 22 families in his Comales besides the above four families whose constituent genera he placed in eight families. Thome (1992) included four suborders in his Comales, nine families in Comineae, and Davidia, Camptotheea, Nyssa, Di­ plopanax, Mastixia, and Cornus in Comaceae. In contrast, others adopted a much nar­ rower circumscription for Comales or Comaceae. Takhtajan (1997) included only eight genera in his Comales and placed the other nine genera in seven separate orders. He adopted narrower family circumscriptions and established eight families among the nine genera of Comales. Eyde (1988) limited his Comaceae to only six genera, including the latest addition of Diplopanax formerly placed in Araliaceae (Eyde & Xiang 1990). He was not sure of the affinity of Alangium with the Comales, but con­ sidered the other genera to be remotely related. In recent years rbeL sequences of the Comaceae alliance have been analysed (Xiang et al. 1993 and Xiang & Soltis in press) based on the circumscription of the Comales by Cronquist (1981) with the addition of Diplopanax. These results showed a Coma­ ceous clade that included only some genera; the other genera were scattered among several other clades. The Comaceous clade is quite close to the Hydrangeaceae. To obtain results comparable to those of the rbeL sequences, we studied all the genera of Cronquist's Comales, and selected three outgroup genera from the taxa used for the rbeL analyses. The Comales in this study consist of Alangiaceae, Comaceae, Garryaceae, and Nyssaceae (Cronquist 1981). Diplopanax is provisionally included in the Comaceae following Eyde & Xiang (1990). The infrageneric subdivision of the studied genera and species circumscription follows Bloembergen (1939) for Alangium with revi­ sions of nomenclature by Eyde (1968), Murrell (1993) for Cornus, Matthew (1976) for Mastixia, and Dahling (1978) for Garrya. Besides these revisions, we follow Fang (1983) for Chinese Alangium, Ricket (1945) for North American Cornus, Soong (1990) for Chinese Mastixia, Aueuba, and Helwingia, Hu (1990) for Cornus s.l. and Torieellia, Burckhalter (1992) for North American Nyssa, Fang (1983) for Asian Nyssa, and Dudley & Santamour (1994) for Cornus angustata.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access 46 IAWA Journal, Vol. 19 (1), 1998

The genus Cornus has been regarded as a single genus in the broad sense or has been subdivided into several genera, and there is as yet no consensus (Murrell1993). In this study we employed the broad genus concept following Eyde (1987). This finds at least some support in the rather uniform wood structure within Cornus S.l. The systematic position of Alangium grisolleoides within Alangiaceae is in dis­ pute. Eyde (1968) included it in sect. Constigma mainly based on fruit structure, but later he doubted his former decision because pollen morphology and wood anatomy suggested a different section (Eyde 1972). After comparing available morphological and chemical characters, Eyde (1988) was still uncertain about its placement within Alangium. According to our observation, wood of this species fits closely with that of sect. Marlea, not only in the type of perforation p1ates, but also in vessel element length and occurrence of paratracheal parenchyma. We therefore treat this species in sect. Marlea. Wood anatomy of the Comales has so far been studied independently within some families, such as Comaceae (Adams 1949), Nyssaceae (Titman 1949), and Garryaceae (Moseley & Beeks 1955), or broadly but using a limited number of twig sampies (Li & Chao 1954). The results ofthese studies were mainly interpreted using the classical Baileyan evolutionary trends of decreasing tracheary element length in advanced taxa, and of correlated changes in other wood anatomical features. Thus taxa with Ion ger andnarrowervessel elements having scalariformperforations with numerous bars were considered primitive (cf. Adams 1949). Majorregional studies of Comales include Kanehira (1926) and Yamabayashi (1938) for Japan and Korea, Tang (1936) forChina, Suzuki et al. (1991) for Nepal, Purkayastha & Bahadur (1977) for India, Moll & Janssonius (1914) and Philipson et al. (1980) for the Malesian region, Patel (1973) for New Zealand, and Sudworth & Mell (1911) and Panshin & De Zeeuw (1980) for North America. These studies are descriptive and do not include systematic analyses. In recent years studies of ecological trends in wood structure have revealed that quantitative features such as vessel element length or vessel diameter are signifi­ cantly correlated with latitude, altitude, or tree size (Baas 1973; Van der Graaff & Baas 1974; Van den Oever et al. 1981; Noshiro et al. 1994, 1995) and that the classi­ cal Baileyan trends are not directly applicable as evolutionary yardsticks for specific taxa. Wheeler & Baas (1991) and Baas & Wheeler (1996) demonstrated the general validity of the Baileyan trends, but also emphasised the rampant occurrence of parallelisms and less so of reversals in accordance with ecological adaptation. As Bailey (1957) hirnself pointed out, the trends in tracheary elements are valid in angiosperms as a whole, but not always applicable when dealing with specific taxa because of deviations from the general trends. Moreover, in cladistic phylogenetic systematics outgroup comparison is used to assess the directionality of character evolution. Character transformations within a group should be evaluated in comparison with the character states of the closest si ster group, not on the basis of general trends in wood evolution. Against this background we embarked on a wood anatomical study covering all the groups of the Comales in order to help determine the relationships within the order and its allies. We aimed at

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 47 comprehensive coverage of the systematic and distributional ranges, especially of wide ranging taxa such as Cornus, in order to grasp total variation and ecological trends in wood structure. This is necessary to build a datamatrix with meaningful character states for the cladistic analysis and to construct a reliable rnicroscopic iden­ tification key.

MATERIALS AND METHODS

We studied 287 specimens belonging to 110 species and 17 genera, mostly obtained from various institutional wood collections (Stern 1978). The list of materials with their locality and specimen numbers is given in the appendix. Whenever available, three specimens were studied per species, and more were included for species distrib­ uted in more than one region for better coverage of their total variation. For some specimens, only sections could be obtained and macerations were not studied. For a limited number of species, only branchwood specimens were available. These spe­ cies were studied mainly for qualitative characters and are not included in the eco­ logical and phylogenetic analyses. Species of Cornus subg. Arctocrania form only a limited amount of secondary xylem in rhizomes and current year branches. For Kaliphora, only branchwood material from a herbarium specimen was available. For these two taxa, only descriptions are given because their wood is not comparable with the mature secondary xylem of the other taxa. Wood blocks were sectioned and macerated according to standard techniques for light microscopy. Quantitative characters of vessels in cross sections were measured with an image analysing system. One to three images (either 0.64 x 0.48 mm, 1.28 x 0.96 mm, or 3.2 x 2.4 mm depending on vessel size and frequency) each with 30-200 vessels were studied for most specimens, and at least 30 vessels per species were measured for specimens with very low vessel frequency. Vessel element length and fibre length were measured from macerations in 30 elements per sampie. Bar number was counted from radial sections and macerations in 30 elements. Vessel grouping is expressed by the index of Carlquist (1988) where the total number of vessels is divid­ ed by the total number of vessel groups. The terminology of wood features follows the IAWA list (1989). The phylogenetic analysis was conducted using PAUP 3.1.1 (Swofford 1993). Search options are describted in the phylogeny section.

DESCRIPTIONS

The main characters of each species are summarised in Table 2 with ranges of means for the quantitative characters. Qualitative features do not show much variation within species, sections or genera except for growth rings and porosity. Quantitative features occasionally differ greatly within species and supraspecific taxa up to the genus level according to their provenance or systematic affinity. For quantitative characters only ranges of means for each specimen are given, and no extreme va1ues are presented. Measurements of branchwood are given in Table 2, but are not considered in the fol­ lowing descriptions. (text continued on page 53)

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Table 2. Wood anatomical characters of the species studied. For legends, see page 51. I~

Characters Species 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Alangium sec!. Alangium longiflorum ± D 1.65-2.02 9-19 55-75 R 530-830 M - 1340-1660 AL 1-3 A Pra salvifolium D 1.80--2.22 13-28 59-93 R 350--680 M - 1010--1880 AL 1-4 A Pra Alangium sec!. Constigma havilandii D 1.44--1.67 12-18 94-99 8-8 R - 1420--1680 - M - 2300--2540 AL 1-5 A Pra javanicum D 1.34-2.32 9-20 80--150 11-16 R 1310-2070 M 2600-3120 AL 1-8 A Pra Sra nobile D 1.61 13 124 7 R 1370 M - 2880 AL 1-5 A Pra Sra ridleyi D 1.56-2.45 10-14 123-148 11-14 R - 1340-1830 - M - 2750--3220 AL 1-5 A Pra Alangium sec!. Marlea alpinum + S 1.18-2.23 15-23 68-89 R 460--640 M - 1270-1760 DA SV 1-4 A PrDra chinense ± S 1.11-1.49 6-30 70-179 R 420--740 M - 1350-1930 A V 1-8 A Pra Dra grif.fithii ± D 1.59-1.91 8-11 114--116 R 690--890 M - 1500-1710 AL S* 1-5 A Pr grisolleoides D 1.83 9 79 R 900 M - 2040 AL S 1-3 A Pra kurzii ± D 1.06--1.65 6-10 95-137 R 490-750 M - 1410-1890 AL SV 1-6 A Pra platanifolium + SR 1.12-1.14 33-49 41-47 R 420-450 M 960-1100 D SV 1-7 A PraDr premnifolium + S 1.09-1.21 12-15 86--88 R 500--530 M - 1160--1570 A V 1-6 A Pra Dr rotundifolium + S 1.25-1.32 3-6 127-133 R 530--610 M - 1540--1830 D SV 1-5 A Alangium sec!. Rhytidandra villosum ± D 1.49-2.23 10--36 52-94 R 680-990 M - 1430--1820 AL 1-5 A Pr Aralidium pinnatifidum D 1.09-1.42 12-23 84--93 25-31 R - 1210-1240 - M - + 1820-2090 S 5-14 B Aucuba chinensis ± D 1.02-1.23 62-80 28-34 27-29 B ++ 1020--1020 + M - ± 1710--1710 S 1-6 B ...... himalaica + D 1.06 41 36 48 B ++ 850 + M - ± 1110 D S 1,6-10 B Downloaded fromBrill.com10/10/2021 08:45:33PM ~ japonica + D 1.02-1.06 54--129 26-32 52-70 B ++ 900-1020 + M ± 1220--1510 S 1,3-10 B ;J> Comus subg. Afrocrania '-< volkensii D 1.67-1.73 21-43 70-102 40-47 B - 1490-1700 - D 2030-2170 DA 1-5 A e0 Comus subg. Arctocrania 3 canadensis** -/± D 1.27-1.7 550-610 26 21-27 B 400-660 D D ? Cornus subg. Cornus - chinensis + D 1.02-1.03 49-65 50-56 39-52 B * 890-1430 D 1200--2050 A 1-4 A ~ mas + D 1.01-1.06 56-85 36-48 27-29 B 930-990 D 1250-1610 A 1-4 A ..... 'Ci officinalis + D 1.01-1.02 58-102 39-40 24-34 B - 620--1290 - D 1530-1740 A 1-4 A ~ sessilis + D 1.03-1.06 82-115 42-48 26-28 B * 730-840 D 1000-1060 A 1-4 A ..... Cornus subg. Cynoxylon ---..... florida D 1.03-1.04 45-80 53-59 33-40 B -* 940-1200 - D 1580--1750 A 1-7 A 'Ci via freeaccess + 'Ci nuttallii + D 1.04-1.08 57-69 48-65 31-37 B - 1020-1220 - D 1290-1710 A 1-4 A 00 subg. Z Comus Discocrania 0 disciflora D 1.02-1.07 28-49 67-87 27-33 B - 1210-1530 - D 1770-2280 A 1-5 A PrDr ::r'" Cornus subg. Kraniopsis a· alba + DS 1.04-1.07 96--107 39-41 28-35 B - 710-1030 - D 980-1320 D 1-3 A alsophila** + D 1.02 98 36 I B I D I D 1-3 A Ro amomum + D 1.02-1.06 75-173 36--50 20-33 B 800--960 D 1130-1430 A 1-5 A I:lj III angustata + D 1.02 45 48 51 B * 1520 D 2270 A 1-6 A Pr III asperifolia + D 1.04 70 40 22 B 1000 D 1280 D 1-4 A '" bretschneideri + D 1.01 38 63 20 B 1100 D 1570 D 1-9 A coreana D 1.05 30 64 15 B 850 D 1500 A 1-5 A CZl + '< darvasica** + D 1.02 116 32 I B I D I A 1-4 A '" drummondi + D 1.04-1.05 102-102 39-46 21-25 B 910-980 D 970-1570 A 1-3 A 8<> excelsa + D 1.06 72 44 24 B 1220 D 1870 A 1-5 A ~. glabrata** + D 1.06 160 37 I B I D I A 1-3 A (') macrophylla + D 1.00-1.06 44-76 48-73 23-44 B -* 970-1260 D 1570-2000 A 1-5 A ~ meyeri** + D 1 64 32 I B I D I A 1-4 A 0 0 mombergii + D 1 33 56 33 B 1000 D 1540 A 1-5 A 0- occidentalis + DS 1.02-1.05 57-86 46--58 32-40 B 560-980 D 960-1270 A 1-4 A § parviflora + D 1.00-1.01 46-57 50-54 50-71 B -* 1210-1480 - D 1600-1960 A 1-8 A ~ peruviana D 1.01-1.02 27-40 57-71 39-39 B 1450-1450 D 2190-2190 A 1-6 A 0 1.01 44 30 B 710 D 1250 A 1-4 A 8 poliophylla + S 58 '< purpusii D 1.02-1.06 112-130 39-41 32-34 B 890-980 D 1140-1440 A 1-2 A + 0 racemosa + D 1.02-1.03 90-170 31-40 20-25 B 700--940 D 1040-1270 A 1-3 A ...., rugosa + D 1.03-1.04 83-208 31-48 26--32 B - 820-1090 - D 1190-1410 A 1-3 A (') 0 sanguinea + D 1.03-1.05 71-81 43-45 24-28 B -* 730-870 D 1130-1300 A 1-5 A ::1 subsp. australis* * + D 1.02 78 34 I B I D I A 1-3 A III (') schinderi** + D 1.02 93 34 I B I D I D 1-2 A CD stolonifera + D 1.ü1-1.05 93-179 34-41 32-35 B -* 690-890 D 900-1050 A 1-2 A ~ stricta + D 1.02-1.03 69-85 45-62 28-35 B * 910-1130 D 1510-1710 A 1-6 A

Downloaded fromBrill.com10/10/2021 08:45:33PM walteri + D 1.01-1.03 31-52 57-71 17-35 B -* 1030-1070 - D 1460-1610 A 1-4 A wilsoniana + D 1.00-1.02 24-40 66--67 14-15 B - 1040-1130 - D 1600-1850 A 1-4 A Cornus subg. Mesomora alternifolia + D 1.14-1.32 85-108 41-52 41-44 B 820-910 D 1200-1290 D 1-4 A controversa + D 1.06-1.24 29-52 60-72 31-53 B 880-1460 - D 1200-2060 DA 1-5 A Cornus subg. Syncarpea capitata + D 1.03-1.08 48-138 33-54 44-50 B -* 1340-1560 D 1910-2020 DA 1-6 A jerruginea + D 1.06 45 50 47 B 1600 D 2360 A 1-6 A hongkongensis + D 1.01-1.03 36-79 45-62 40-60 B -* 1080-1420 - D 1780-2350 A 1-6 A Pr kousa + D 1.00--1.05 34-66 46--59 39-51 B -* 1020-1160 - D 1710-2030 A 1-8 A Pr tonkinensis + D 1.01 70 67 52 B 1570 D 2060 A 1-5 A Cornus subg. Yinquania via freeaccess oblonga D 1.02-1.03 49-90 51-59 20-36 B -* 940-1230 D 1330-2060 A 1-4 A 1.1:>- + \0 (Table 2 continued) Ulo Characters Species 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 2\

Corokia buddleoides + D 1.01 233 23 21 B ++ 720 + M - + 840 1-3 B collenettei D 1.08-1.23 153-216 29-36 17-22 B ++ 700-720 M - + 840-920 1-4 BtA cotoneaster + D 1.01 452 13 16 B ++ 370 + M - + 470 1-3 B macrocarpa + D I 249 17 16 ß ++ 630 + M - ± 760 1-2 B whiteana ± D 1.02-1.09 208-247 18-18 22-23 B ++ 610-690 + M - + 920-1060 1-4 B Curtisia dentata D 1.00-1.00 37-52 48-55 26-39 ß - 1300-1990 - D 2210-2560 A 1-4 A Pr Diplopanax stachyanthus + D 1.05 117 49 34 B-R - 1350 D 1770 DA 1-4 A Griselinia littoralis ± D 1.02 209 34 16 R 800 D ± 1010 A 1-6 B Pr lucida ± D 1.00-1.01 26-35 61-70 14-16 R - 970-1220 - D ± 1260-1540 A 1-6 B Pr ruscifolia ± D 1.03-1.12 200-245 23-39 10-13 R 720-770 D ± 910-1170 A 1-10 B Pr Helwingia himalaica + D 1.06 103 33 27 R M - + D* S 1-11 B Sra japonica + D 1.18-1.24 84-183 24-33 17-41 R 760-940 M + 890-1130 D* S 1-4 B Kaliphora madagascariensis** ± 0 1.45-1.77 134-160 27-33 B D S 1-3 B Mastixia subg. Manglesia octandra D 15 89 54 R + 2280 D 4230 A S 1-6 A Mastixia subg. Mastixia (Alternae) arborea D 1.02-1.05 24-27 117-119 42-51 R + 1970-2330 - D 2990-3330 A S* 1-6 A Pr

Downloaded fromBrill.com10/10/2021 08:45:33PM cuspidata D 1.00-1.01 28-29 90-106 58-59 R + 2460-2620 - D 3420-3450 A S 1-5 A + Pr ~ macrophylla D 1.00-1.01 34-43 73-88 60-66 R + 2030-2310 - D 3070-3190 A S 1-6 A ;J> pentandra D 1.00-1.04 26-43 77-100 45-60 R + 1980-2400 - D 3340-3570 A S 1-6 A Pr ...... o tetrandra D 1.00-1.02 34-47 80-85 47-58 R + 2090-2180 - D 3410-3450 A S 1-6 A <= tetrapetala D I 40 70 48 R + 1640 D 2180 A S 1-4 A 8 Mastixia subg. Mastixia (Oppositae) ? eugenioides D 1.03 45 78 52 R + 2760 D 3830 A S 1-4 A kaniensis D 1.05 26 93 44 R + 1920 D 2930 A S 1-13 A ~ rostrata D 1.üI-1.05 37-46 61-88 38-49 R + 1460-2330 - D 2140-3460 A S 1-8 A ± Pr -'Cl trichotoma D 1.00-1.03 24-45 90-109 45-54 R + 1730-2430 D 2650-3850 A S 1-8 A ± Pr Dr MelanophyUa y capuronii D 1.75 23 88 m R 1070 M ± 1840 S 1,4-8 B 'Cl via freeaccess -'Cl crenata D 2.13 27 92 m R 1170 M + 2040 S 1,2,4-6 B 00 Torieellia oz arguta + R 1.62 84 39 R ++ 480 M + 1140 S 1-4 B tiliifolia + S 1.71-2.04 16-50 73-127 R ++ 560-570 M + 1170--1460 S 1-6 B [ Garrya subg. F adyenia Re fadyenii + o 1.00-1.00 73-117 32-40 6-7 B ++ 650--710 o + 1130--1230 A S 1-6 B l:Jj glaberrima + o 1.01 216 22 5 B ++ 530 o + 810 A S 1-6 B E5 laurifolia + o 1.00-1.02 48-142 34-53 6-8 B ++ 630-770 o + 1050-1490 A S 1-8 B CJ> longifolia + o 1.00--1.01 43-69 47-53 7-7 B ++ 780-830 o ± 1550-1670 A S 1-10 B ova ta + o 163 26 4 B ++ 500 o + 810 o S 1-6 B CI) wrightii + o 1.00--1.01 193-332 16-23 3-4 B ++ 400--520 o + 590-720 DA S 1-8 B '< Garrya subg. Garrya buxifolia + S 1.00--1.07 132-212 24-27 3-5 B ++ 420-450 o + 520-760 DA S 1-10 B ~ d-. elliptiea + o 1.00--1.05 135-220 23-32 4-5 B ++ 530--730 o + 840--1140 DA S 1-11 B () flaveseens + o 1.00-1.01 157-304 23-27 4-5 B ++ 450-560 o + 730--830 DA S 1-11 B ~ fremontii + o 1.00-1.00 142-152 27-30 4-5 B ++ 450-590 o + 600-960 o S 1-10 B o veatehii + o 1.üI-1.05 210-362 21-25 3-4 B ++ 510-540 o + 700-800 DA S 1-10 B 8. Camptotheea § acuminata + o 1.05-1.18 63-80 48-60 17-21 B + 760-1330 o 1280-2010 o 1-3 A Pa Davidia involucrata + o 1.08-1.16 69-100 44-60 60-91 B - 1020-1590 - o 1500--2210 DA - 1-3 A J Nyssa ....,o aquatica + o 1.21-1.88 41-138 36-61 29-40 B + 940-1240 o 1550-1990 o S' 1-2 A Pa () biflora + o 1.13-1.19 26-123 43-54 34-44 B + 860--1140 o 1320-2070 o 1-3 A Pa javanica ± o 1.11-1.34 11-23 77-109 36-50 B + 1650-2080 - o ± 1790--2900 o 1-6 A Pa ogeehe o 1.17-1.51 35-91 53-65 20-23 B + 980--1020 1530--1820 S 1-3 A ~ + o o Pa 0> sinensis + o 1.14-1.22 51-94 56-66 35-44 B + 1240-1340 o ± 2020-2130 o S' 1-3 A Pa f'i sylvatica + o 1.19-1.32 72-150 44-57 25-39 B + 1020--1070 - o 1110--1780 o S 1-3 A Pa Downloaded fromBrill.com10/10/2021 08:45:33PM ursina + o 1.18 96 44 39 B + 850 o 1780 o S 1-2 A Pa

•• =branchwood or rhizome (c. eanadensis): 1= no measurement. - 1: Growth rings: + =present, - = absent. - 2: Porosity: 0 = diffuse, S = semi-ring-porous, R = ring-porous. -

3: Vessel grouping index. - 4: Vessel frequency (I mm 2). - 5: Tangential vessel diameter (~m).- 6: Perforation plate (number of bars): - = simple; m = mixed simple and scalariform. -7: Vessel-ray pits: B = bordered, R = with reduced borders. - 8: Helical thickenings: ++ = throughout the body of vessel elements, + = vessel element ends, - = absent; • = occasionally fine helical thickenings at vessel element ends. - 9: Vessel element length (~).-10: Vascular tracheids: + = present, - = absent. -11: Fibre pits: 0 = distinctly bordered, M = minutely

bordered. - 12: Helical thickenings in fibres: + = present, - = absent. - 13: Septate fibres: + = all, ± = some, - = absent. - 14: Fibre length (~m).- 15: Apotracheal parenchyma: L = irregular tangentiallines, A = diffuse-in-aggregates, 0 = diffuse; • = rare. - 16: Paratracheal parenchyma: S = scanty, V = vasicentric; * = rare. - 17: Ray width (cells). - 18: Ray type: A = body mostly of procumbent cells, B = body includes square/upright cells. -19: Axial canals: + = present, - = absent. - 20: Crystals: P = prismatic crystals, 0 = druses; r = rays, a = axial parenchyma. - 21: Silica grains: S = present; r = rays, a = axial parenchyma. via freeaccess U\ 52 IAWA Journal, Vol. 19 (l), 1998

Fig. 1-3. Alangium chinense (TWTw 7420) - I: TS, semi-ring-porous wood with aliform to confluent latewood parenchyma. - 2: TLS, heterocellular rays. - 3: RLS, simple perforation and vessel-ray pits with reduced borders. - Fig. 4-6. Alangium javanicum (BZFw 30875) - 4: TS, diffuse-porous wood with vessel multiples and parenchyma in irregular lines. - 5: TLS, heterocellular rays and dense intervessel pits. - 6: prismatic crystals in ray cells. - Fig. 7 & 8. Alangium nobile (FHOw 12683) - 7: silica grains in ray cells. - 8: scalariform perforation. - Scale bars =250 J.lm in Fig. 1, 4; 100 J.lm in Fig. 2,5; 50 J.lm in Fig. 3; 12 J.lm in Fig. 6; 6 J.lm in Fig. 7; 30 J.lm in Fig. 8.

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ALANGIACEAE

Alangium - Fig. 1-8 Growth rings usually absent in sect. Alangium, Constigma (Fig. 4), and Rhytidandra, except for two specimens of A. longiflorum (FHOw 12631, FPAw 23717) and one specimen of A. villosum (SJRw 37481); usually present in sect. Marlea (Fig. 1), but indistinct or absent in A. griffithii, A. grisolleoides, and A. kurzii; growth rings marked by differences in vessel diameter between latewood and earlywood and marginal pa­ renchyma. Wood diffuse-porous in sect. Alangium, Constigma (Fig. 4), and Rhyti­ dandra; semi-ring-porous in sect. Marlea (Fig. 1) except for one sampie of A. chinense (FHOw 5067), A. griffithii (all sampies), A. grisolleoides, and A. kurzii (CAFw 3464, FHOw 12622). - Vessels solitary, or in radial multiples or occasional clusters of 2-6 (-10); vessel grouping index 1.06-2.45, low (mostly lower than 1.5) in semi-ring­ porous species in sect. Marlea; 3-49 Imm2; 41-180 11m in tangential diameter; mostly round in outline, occasionally weakly angular in smaller vessels of semi-ring-porous species. Perforations simple in sect. Alangium, Marlea (Fig. 3), and Rhytidandra; scalariform with 7-16 bars in sect. Constigma (Fig. 8). Intervessel pits alternate and dense, often polygonal, 4-14 11m in horizontal diameter (Fig. 5); without vestures. Vessel-ray pits with reduced borders (Fig. 3), occasionally unilaterally compound, round to horizontal or vertical, 3-12 11m in horizontal diameter. Helical thickenings absent. Vessel element length 360-990 11m in sect. Alangium, Marlea, and Rhytidan­ dra; 1310-2070 11m in sect. Constigma. Tyloses usually present, often conspicuous. - Fibres with minutely bordered pits in sect. Alangium, Marlea, and Rhytidandra, or with simple pits and slit-like apertures in sect. Constigma; confined to the radial walls; walls thin to thick, 2.5-6 11m, in sect. Alangium, Marlea, and Rhytidandra, very thick, 8-16 11m, with almost closed lumina in sect. Constigma (Fig. 4). Fibre length 960- 2040 11m in sect. Alangium, Marlea, and Rhytidandra; 2300-3220 11m in sect. Con­ stigma. F IV ratio 1.5-3.9. - Axial parenchyma diffuse-in-aggregates to in many ir­ regular lines in sect. Alangium, Constigma (Fig. 4), and Rhytidandra; in sect. Marlea, diffuse to diffuse-in-aggregates in temperate species, and diffuse-in-aggregates to in irregular lines in tropical species, with additional scanty paratrachtal to vasicentric parenchyma in the whole seetion, occasionally aliform to confluent in latewood of temperate species (Fig. 1); usually in marginal or seemingly marginal bands in sect. Marlea. Parenchyma strands 4-12(-16) cells long in sect. Alangium, Marlea, and Rhytidandra; 8-28 cells long in sect. Constigma. - Rays heterocellular, 1-5(-8) cells wide, up to 1.4(-2.5) mrn tall (Fig. 2, 5); composed of procumbent body cells and 2-9 marginal rows of upright or square cells; sheath cells rare in A. javanicum and A. platanifolium; uniseriate rays consisting solely of square or upright cells. - Prismatic crystals usually conspicuous in both ray cells (Fig. 6) and axial parenchyma, but rare in the latter in sect. Rhytidandra; druses occasionally present in ray cells and axial parenchyma of temperate species of sect. Marlea; silica grains infrequent in ray cells and axial parenchyma of A. javanicum (FHOw 11942, FHOw 12681, SANw 98859) and A. nobile (FHOw 12683) in sect. Constigma (Fig. 7).

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Fig. 9-11. Aralidium pinnatifidum (9, 10: SJRw 16054, 11: MADw 27358) - 9: TS, diffuse-porous wood and paratracheal parenchyma. - 10: TLS, large multiseriate rays and absence of uniseriate ones. - 11: RLS, scalariform perforations and upright or square cells in the body of a multiseriate ray. - Fig. 12-14. Aucubajaponica (TWTw 5688) - 12: TS, diffuse-porous wood with small vessels. - 13 : TLS, large mul­ tiseriate rays with some sheath cells and short uniseriate rays. - 14: RLS, scalariform perforations and helical thickenings in vessels and vascular tracheids. - Scale bars = 250 J.Im in Fig. 9, 10, 12.13; 100 J.Im in Fig. 11; 50 J.Im in Fig. 14.

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CORNACEAE

Aralidium - Fig. 9-11 Growth rings absent (Fig. 9). Wood diffuse-porous, with a tendency to a radial vessel pattern dissected by wide rays (Fig. 9). - Vessels solitary or in radial multi­ ples or oblique (to tangential) clusters of 2-4, vessel grouping index 1.09-1.42, 12-

23 I mm2, 84-93 lJ1ll in tangential diameter, angular in outline. Perforations scalari­ form with 25-31 bars (Fig. 11). Intervessel pits opposite-scalariform, 8-10 J.lm in horizontal diameter; without vestures. Vessel-ray pits with reduced borders, round to horizontal, 4-40 J.lm in horizontal diameter (Fig. 11). Helical thickenings absent. Vessel element length 1210-1240 1J1ll. - Fibres with minutely bordered pits mostly in radial walls; all septate (Fig. 10); walls thin to thick, 5-12 J.lm thick; 1820-2090 IJ1lll0ng. F/V ratio 1.5-1.7. - Axial parenchyma scanty paratracheal (Fig. 9); 4-12 cells per strand. - Rays heterocellular, 4-12 cells wide, very rarely uniseriate, up to 5.5 mm tall (Fig. 10); often with sheath cells; rays composed mainly of procumbent body cells, but including square or upright cells (Fig. 11), with 1 or 2 marginal rows of upright cells. - Crystals absent.

Aucuba - Fig. 12-14 Growth rings distinct (Fig. 12), marked by one row of slightly larger earlywood vessels aligned discontinuously along growth rings and radially flattened fibres in the latewood; indistinct in one sampie of A. chinensis (CAFw 151). Wood diffuse-po­ rous, with a tendency to a radial vessel pattern dissected by wide rays (Fig. 12). - Vessels mostly solitary or rarely in multiples or clusters of 2 or 3; vessel grouping index 1.02-1.23; 41-129/mm2; 26-36 J.lm in tangential diameter, angular in outline. Perforations scalariform with 27-70 bars (Fig. 14); 27-29 bars in A. chinensis. Intervessel pits rare, opposite( -scalariform), 6(-20) J.lm in horizontal diameter; with­ out vestures. Vessel-ray pits with distinct borders, round (to horizontal) and dense, 4-15 J.lm in horizontal diameter (Fig. 14). Helical thickenings distinct throughout ves­ seI elements (Fig. 14). Vessel element length 850-1020 J.lm. Vascular tracheids with distinct helical thickenings around vessels (Fig. 14). - Fibres with minutely border­ ed pits in radial walls, occasionally septate, walls thin to thick, 6 J.lm; 1110-1710 lJ1ll long. F IV ratio 1.3-1.7. - Axial parenchyma scanty paratracheal, rarely diffuse. - Rays heterocellular; of two distinct sizes in A. himalaica and A. japonica, 1 and 3-10 cells wide, up to 6.9 mm tall (Fig. 13); and 1-6 cells wide, up to 4.5 mm tall in A. chinensis; rays consisting of a mixture of procumbent, square, and upright body cells, with 1-5 marginal rows of upright cells; with sheath cells. - Crystals absent.

Cornus - Fig. 15-27 Growth rings usually present (Fig. 15, 19) and marked by slight differences in vessel size between latewood and earlywood and radially flattened fibres at the end of growth rings; absent in three tropical species, C. disciflora, C. peruviana and C. volkensii (Fig. 22, 25). Wood diffuse-porous, with a tendency to semi-ring-porosity in C. alba (Tw 43798), C. occidentalis (PL 1535), and C. poliophylla (PL 1537) of

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Fig. 15-17. Comus macrophylla (TWTw 13509) - 15: TS, diffuse-porous wood with exclusively solitary vessels and diffuse-in-aggregates parenchyma. - 16: TLS, heterocellular rays. - 17: RLS, scalariform perforation and vessel-ray pits. - Fig. 18. Comus kousa (TWTw 13511), resin cast of avesseI element tai! with fine helical thickenings. - Fig. 19-21. Comus controversa (TWTw 13356) - 19: TS, diffuse­ porous wood with some vessel multiples. - 20: TLS, opposite-scalariform intervessel pits. - 21: RLS, small vessel-ray pits with distinct borders. - Scale bars = 250 11m in Fig. 15, 19; 100 11m in Fig. 16; 50 11m in Fig. 17, 20, 21 ; 12 11m in Fig. 18.

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Fig. 22-24. Cornus disciflora (MADw 24710) - 22: TS. diffuse-porous wood with dense, exclusively solitary vessels. - 23: TLS, heterocellular rays. - 24: RLS, scalariform perforation and vessel-ray pits. - Fig. 25 & 26. Cornus volkensii (SJRw 27557) - 25: TS, diffuse-porous wood with frequent vessel multi­ ples. - 26: TLS, heterocellular rays with some sheath cells. - Fig. 27. Cornus canadensis (Hatta s.n., 6 Aug. '79), TS, diffuse-porous wood with den se narrow vessels. - Scale bars = 250 f.lm in Fig. 22, 23, 25; 100 f.IID in Fig. 26, 27; 50 f.lm in Fig. 24.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access 58 IAWA Journal, Vol. 19 (1), 1998 subg. Kraniopsis; vessel diameter gradually decreasing towards the growth ring bound­ ary in species with distinct growth rings. - Vessels mostly solitary or rarely in ra­ dial multiples of 2 or 3, with avessei grouping index of 1.00-1.08 in subg. Cornus, Cynoxylon, Discocrania (Fig. 22), Kraniopsis (Fig. 15), Syncarpea, and Yinquania; solitary or in radial multiples (rarely oblique clusters) of2-4, and with avessei grouping index of (1.06-)1.14-1.73 in subg. Afrocrania (Fig. 25) and Mesomora (Fig. 19); 21-208/mm2; 31-102 flm in tangential diameter, relatively wide (over 60 flm) in tropical species; round-angular to angular in outline. Perforations scalariform with 14-71 bars, usually more than 40 in subg. Afrocrania and Syncarpea (Fig.17, 21,24). Intervessel pits opposite-scalariform, 6-10 flm in horizontal diameter, dense in subg. Afrocrania and Mesomora (Fig. 20); without vestures. Vessel-ray pits with distinct borders (Fig. 17,21,24), occasionally unilaterally compound; round and dense, op­ posite to alternate, rarely horizontal, 4-6(-15) flill in diameter. Helical thickenings usually absent; fine thickenings occasionally occur in vessel element tails of some specimens of subg. Cornus, Cynoxylon, Kraniopsis, Syncarpea (Fig. 18), and Yinqua­ nia. Vessel element length 560-1710 flm, longest (mostly over 1400 flm) in tropical species. - Fibres with distinctly bordered pits in both radial and tangential walls, walls thin to thick, 3-8 flm; 900-2360 flm long, longest (mostly over 2000 flm) in tropical species. F IV ratio 1.1-1.8(-2.5). - Axial parenchyma diffuse or diffuse-in­ aggregates (Fig. 15, 19,22,25),4-12(-15) cells per strand. - Rays heterocellular, 1-6(-9) cells wide, up to 2 mm tall, composed of procumbent body cells with 1-5 marginal rows of upright or square cells; rarely with sheath cells (Fig. 16,20,23,26). - Crystals rare; prismatic crystals in C. peruviana (MADw 42286), together with druses in C. angustata (CAFw 13429), C. hongkongensis (CAFw 15788), and C. kousa (CAFw 9781). Cornus canadensis (secondary xylem limited to rhizomes and current-year shoots) (Fig. 27): Growth rings indistinct. Wood diffuse-porous. Vessels solitary or in multi­ ples of 2; vessel grouping index 1.27-1.7; 153-453/mm2, 21-27 flm in tangential diameter, angular in outline. Perforations scalariform with about 26 bars. Interves­ seI pits opposite-scalariform; without vestures. Fibres with distinctly bordered pits in radial walls. Axial parenchyma diffuse. Rays uniseriate and entirely composed of upright cells.

Corokia - Fig. 28-30 Growth rings absent in C. collenettei and distinct or indistinct in other species (Fig. 28) rnarked by smaller vessels and radially flattened fibres at the end of growth rings. Wood diffuse-porous or with a tendency to semi-ring-porosity in C. buddleoi­ des (Fig. 28) and C. cotoneaster. - Vessels mostly solitary or occasionally in multi­ ples of2 or 3; vessel grouping index 1.00-1.09(-1.23); 153-453/mm2, 13-36 flm in tangential diameter, round-angular in outline. Perforations scalariform with 16-23 bars (Fig. 30). Intervessel pits opposite-alternate, 4-6 flm in horizontal diameter; without vestures. Vessel-ray pits with distinct borders, round (to horizontal) and crowd­ ed, opposite to alternate, 4-6(-20) flm in horizontal diameter (Fig. 30). Fine helical thickenings present throughout vessel elements. Vessel element length 370-720 flill.

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Fig. 28-30. Corokia buddleoides (SJRw 25438) - 28: TS, diffuse-porous wood with small solitary ves­ sels. - 29: TLS, heterocellular rays. - 30: RLS, scalariform perforations and dense vessel-ray pits. - Fig. 31-34. Curtisia dentata (MADw 16804) - 31: TS, diffuse-porous wood with exclusively solitary vessels and diffuse parenchyma. - 32: TLS, heterocellular rays. - 33: RLS, prismatic crystals and vessel­ ray pits with distinct borders. - 34: RLS, scalariform perforation. - Scale bars =250 11m in Fig. 28, 31; 100 11m in Fig. 29, 32; 50 11m in Fig. 30, 33, 34.

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Vascular tracheids present except in C. collenettei; with distinct helical thickenings in C. cotoneaster and C. macrocarpa. Fibres with minutely bordered pits in radial walls; all septate; walls thin to thick, 6-8 !lIll ; 470-1060 J..lm long. F IV ratio 1.2-1.5. - Axial parenchyma absent (Fig. 28). - Rays heterocellular, 1-4 cells wide, up to 0.9 mm tall (Fig. 29), rays composed of procumbent and square body cells with 1-7 marginal rows of upright cells, with occasional sheath cells and perforated ray cells with about 10 bars in perforations in C. collenettei (SJRw 37337). - Crystals absent.

Curtisia - Fig. 31-34 Growth rings absent (Fig. 31). Wood diffuse-porous (Fig. 31). - Vessels exclu­ sively solitary, very rarely in pairs (Fig. 31); vessel grouping index 1.00-1.01,37-521 mm2, 48-55 J..lm in tangential diameter, round-angular in outline. Perforations sca­ lariform with 26-39 bars (Fig. 34). Intervessel pits opposite(-scalariform), 4-6 J..lm in horizontal diameter; without vestures. Vessel-ray pits with distinct borders, round to horizontal, opposite to scalariform, 4-6(-20) !lIll in diameter (Fig. 33). Helical thickenings absent. - Vessel element length 1300-1990 J..lm. - Fibres with distinctly bordered pits in both radial and tangential walls; walls thin to thick, 8-11 J..lm; 2210- 2560 !lIlllong. F/V ratio 1.3-1.7. -Axial parenchyma diffuse to diffuse-in-aggre­ gates (Fig. 31), 6-12 cells per strand. - Rays heterocellular, 1-4 cells wide, up to 2.2 mm tall, composed of procumbent body cells with 1-5(-10) marginal rows of upright or square cells, often with sheath cells (Fig. 32). - Prismatic crystals common in ray cells (Fig. 33).

Diplopanax - Fig. 35-37 Growth rings distinct (Fig. 35), marked by smaller and fewer vessels and radially flattened fibres at the end of growth rings. Wood diffuse-porous (Fig. 35), vessel diameter and frequency decreasing in the latewood. - Vessels mostly solitary or rarely in multiples of 2 or 3, vessel grouping index 1.05, 117 Imm2, 49 J..lm in tan­ gential diameter, angular in outline (Fig. 35). Perforations scalariform with 34 bars (Fig. 37). Intervessel pits opposite (to scalariform), sparse, 8-15 !lIll in horizontal diameter (Fig. 36); without vestures. Vessel-ray pits with distinct to reduced borders, round to horizontal, opposite to alternate (Fig. 37). Helical thickenings absent. Vessel element length 1350 J..lm. - Fibres with distinctly bordered pits in both radial and tangential walls (Fig. 36, 37); walls thin to thick, 5-6 J..lm; 1770 !lIlllong. F IV ratio 1.3. - Axial parenchyma diffuse to diffuse-in-aggregates (Fig. 35), 4-9 cells per strand. - Rays heterocellular, 1-3(-4) cells wide (Fig. 36), composed of procumbent body cells and 1-12 marginal rows of upright or square cells; sheath cells very rare. Crystals absent.

Griselinia - Fig. 38-40 Growth rings indistinct to distinct (Fig. 38), marked by differences in vessel size between latewood and earlywood. Wood diffuse-porous (Fig. 38), usually with a slight reduction in vessel size in the latewood. - Vessels mostly solitary or occasion­ ally in pairs, vessel grouping index 1.00-1.12, 26-245/mm2, 23-70 J..lm in tangential

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Fig. 35-37. Diplopanax stachyanthus (CAFw 15828) - 35: TS, diffuse-porous wood with dense solitary vessels. - 36: TLS, heterocellular rays. - 37: RLS, scalariform perforations and sparse vessel-ray pits with distinct to reduced borders. - Fig. 38-40. Griselinia lucida (SJRw 50340) - 38: TS, diffuse-porous wood with an indistinct growth ring boundary. - 39: TLS, heterocellular rays with several sheath cells. - 40: RLS, scalariform perforation and vessel-ray pits with reduced borders. - Scale bars = 250 11m in Fig. 35,38; 100 11m in Fig. 36, 39; 50 11m in Fig. 37,40.

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Fig. 41-44. Helwingia japonica (TWTw 14374) - 41: TS, diffuse-porous wood with solitary vessels, showing a tendency to semi-ring-porosity. - 42: TLS, heterocellular rays. - 43 : RLS, round to horizontal vessel-ray pils with reduced borders. - 44: RLS, scalariform perforation. - Fig. 45 - 47. Kaliphora madagascariensis (Bernardi 11058) - 45 : TS, diffuse-porous wood with vessel multiples. - 46: TLS, het­ erocellular rays. - 47: RLS , simple and scalariform perforations and vessel-ray pits with distinct borders. - Fig. 48. He/wingia himalaica (Suzuki et al. 8840197), RLS, silica grains in ray cells. - Scale bars = 250 /Im in Fig. 41 , 45; 100 /Im in Fig. 42, 46; 50 /Im in Fig. 43, 44; 25 /Im in Fig. 47, 48.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 63 diameter, angular in outline. Perforations scalariform with 10-17 bars (Fig. 40). In­ tervessel pits opposite to scalariform, 7-20 11m in horizontal diameter; without vestures. Vessel-ray pits with reduced borders, round to horizontal, opposite to alter­ nate (Fig. 40). Helical thickenings absent. Vessel element length 720-1220 Ilill. Tyloses present only in one sampie of G. littoralis (FPAw 10013). - Fibres with distinctly bordered pits in both radial and tangential walls; occasionally septate, grading into axial parenchyma strands (Fig. 39); walls thin to thick, 4-6 11m thick, 910-1540 11m long. F IV ratio 1.3-1.5. -Axial parenchyma diffuse-in-aggregates, 3-8 cells per strand. - Rays heterocellular, 1-10 cells wide, up to 3 mm in G. lucida (Fig. 39) and G. littoralis, 10 mm or more tall in G. ruscifolia, composed of mostly procumbent body cells, but including square or upright body cells (Fig. 39), with 1-3 marginal rows of upright cells; with occasional sheath cells. - Prismatic crystals usually in body ray cells.

Helwingia - Fig. 41-44, 48 Growth rings distinct (Fig. 41), marked by differences in vessel size between latewood and earlywood and radially flattened fibres in the latewood. Wood diffuse­ porous, with a gradual reduction in vessel diameter towards the growth ring bounda­ ries, occasionally with a tendency to semi-ring-porosity (Fig. 41). - Vessels mostly solitary or in multiples of 2 or 3, vessel grouping index 1.02-1.24, 84-183/mm2, 24- 33 11m in tangential diameter, angular in outline. Perforations scalariform with 17-41 bars (Fig. 44). Intervessel pits opposite to scalariform, 8 Ilill or more in hori­ zontal diameter; without vestures. Vessel-ray pits with reduced borders, round to horizontal, 4-35 Ilill in horizontal diameter (Fig. 43). Helical thickenings absent. Vessel element length 760-940 Ilill. Tyloses occasionally present. - Fibres with mi­ nutely bordered pits in radial walls, all septate (Fig. 42); walls thin to thick, 2.5-5 11m; 890-1130 11m long. F/V ratio 1.1-1.4. - Axial parenchyma scanty paratracheal (Fig. 44),4-8 cells per strand. - Rays heterocellular, 1-11 cells wide, up to 7 mm or more tall, composed of procumbent, square, and upright body cells, with 1-3 mar­ ginal rows of upright cells, with occasiona1 sheath cells (Fig. 42); uniseriate rays composed of upright cells on1y. Silica grains present in ray cells (Fig. 48) and axial parenchyma, and also in septate fibres of H. himalaica (Suzuki et al. 8840197).

Kaliphora (description based on branchwood specimens of c. 4.5 mm in diameter) -Fig.45-47 Growth rings indistinct (Fig. 45), marked by differences in fibre diameter between latewood and earlywood. Wood diffuse-porous with a laxly radial vessel alignment (Fig. 45). - Vessels solitary or in radial multiples of 2-5(-9), vessel grouping index 1.45-1.77, 134-160/mm2, 27-33 11m in tangential diameter, round-angular in out­ line. Perforations simple, rarely scalariform with 1 or 2 bars (Fig. 47). Intervessel pits alternate, round and dense, 4 11m in diameter; without vestures. Vessel-ray pits with distinct borders, dense, alternate to opposite, c. 3 11m in diameter (Fig. 47). Heli­ cal thickenings absent. - Fibres with distinctly bordered pits mostly in radial walls; walls thick to thin, 3.5-5 Ilill. - Axial parenchyma scanty paratracheal and rare. -

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Fig. 49-51. Mastixia cuspidata (SANw 132556) - 49: TS, diffuse-porous wood with dense solitary pores and axial canals. - 50: TLS, heterocellular rays with some sheath cells. - 51: RLS, scalariform perforation and horizontal vessel-ray pits. - Fig. 52-54. Melanophylla capuronii (CTFw 9052) - 52: TS, diffuse­ porous wood with many vessel multiples. - 53: TLS, large heterocellular rays and small uniseriate ones. - 54: RLS, simple perforation and dense vessel-ray pits. - Fig. 55. Melanophylla crenata (CTFw 8879), RLS, simple and scalariform perforations. - Scale bars =250 J.lm in Fig. 49, 52, 53; 100 J.lm in Fig. 50, 51,54.55.

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Rays heterocellular, 1-3 cells wide, composed of procumbent, square, and upright body and marginal cells, with rare sheath cells (Fig. 46). - Crystals absent. Infrequent occurrence of scalariform perforation and frequent occurrence of radial vessel multiples may suggest affinity to another Madagascar taxon, Melanophylla. Septate fibres may be found in mature wood. The distinctly bordered pits in fibre walls disagree with this placement, but they are small in diameter, c. 4 ~m.

Mastixia - Fig. 49-51 Growth rings absent (Fig. 49). Wood diffuse-porous (Fig. 49). - Vessels almost exclusively solitary, rarely in multiples of 2 or 3, vessel grouping index 1.00-1.05, 24-47/mm2 in subg. Mastixia and 15/mm2 in M. octandra (Meijer 6316) in subg. Manglesia, 61-119 11m in tangential diameter, angular in outline. Perforations sca­ lariform with 38-66 bars (Fig. 51). Intervessel pits opposite-scalariform, 8 /lffi or more in horizontal diameter; without vestures. Vessel-ray pits with reduced borders, occasionally unilaterally compound, round to horizontal, 6-50 11m in horizontal diam­ eter (Fig. 51). Distinct helical thickenings present in vessel element tails. Vessel ele­ ment length 1460-2760 /lffi. Tyloses present in one specimen of M. trichotoma (SANw 59252). - Fibres with distinctly bordered pits in both radial and tangential walls; walls thin to thick, 4-11 11m; 2140-3850 /lffi long in subg. Mastixia and 4130 11m in M. octandra. F IV ratio 1.3-1.7 in subg. Mastixia and 1.8 in M. octandra. - Axial parenchyma diffuse-in-aggregates and scanty paratracheal (Fig. 49), 5-12 cells per strand. - Rays 6-12 per mm, heterocellular, 1-8(-13) cells wide, up to 3.6 mm tall, composed of procumbent body cells with 1-5 marginal rows of upright or square cells; often with sheath cells (Fig. 50); uniseriate rays consisting of 1-12 rows of up­ right cells. Axial canals occasionally in long tangentiallines in M. cuspidata (SANw 132556, van Balgooy & van Setten 5471) (Fig. 49), M. rostrata (de Wilde & de Wilde­ Duyfjes 12714, SJRw 31022), andM. trichotoma (de Vogel 1249). - Prismatic crys­ tals occasionally present in body ray cells; druses noted in ray cells of one specimen of M. trichotoma (SANw 59252).

Melanophylla - Fig. 52-55 Growth rings absent (Fig. 52). Wood diffuse-porous (Fig. 52). - Vessels solitary or in radial multiples or clusters of 2-6(-9), vessel grouping index 1.75-2.13, 23-27 Imm2, 88-92 11m in tangential diameter, angular in outline. Perforations mostly sim­ ple with distinct rims or occasionally scalariform with 5-24 bars (Fig. 54, 55). Intervessel pits opposite to scalariform, crowded, 8 /lffi or more in horizontal diam­ eter (Fig. 53); without vestures. Vessel-ray pits with reduced borders, occasionally unilaterally compound, opposite to horizontal and dense (Fig. 54), 8-35 11m in hori­ zontal diameter. Helical thickenings absent. Vessel element length 1070-1170 /lffi. - Fibres with minutely bordered pits in radial walls, mostly septate; walls thick to thin, 6-8 11m; 1840-2040 11m long. F/V ratio 1.7-1.8. - Axial parenchyma scanty paratracheal (Fig. 52), 4-8 cells per strand. - Rays heterocellular, of two distinct sizes, 1-2 and 4-8 cells wide, up to 4.8 mm tall; composed of procumbent and square cells, with occasional sheath cells (Fig. 53). - Crystals absent.

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Fig. 56-58. Garrya elliptica (MADw 45076) - 56: TS , diffuse-porous wood with small solitary vessel. - 57: TLS, heterocellular rays with many sheath cells. - 58: RLS, scalariform perforations and helical thickenings in vessels. - Fig. 59-61. Camptotheca acuminata (SJRw 21463) - 59: TS, diffuse-porous wood with vesse1 multiples. - 60: TLS, heterocellular rays. - 61: RLS, scalariform perforation and vessel­ ray pits. - Scale bars = 250 !Im in Fig. 56, 59; 100 !Im in Fig. 57, 60; 50 !Im in Fig. 58, 61.

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Toricellia - Fig. 71-73 Growth rings distinct (Fig. 71), marked by differences in vesse1 diameter between 1atewood and ear1ywood and radially flattened fibres at the end of growth rings. Wood semi-ring-porous in T. tiliifolia; ring-porous with a discontinuous row of mostly soli­ tary earlywood pores and latewood pores in a radial to diagonal pattern in T. arguta (Fig. 71). - Vessels solitary or in multiples or clusters of 2-8, vessel grouping index

1.62-2.04, vessel frequency 16-84/mm2, 39-127 11m in tangential diameter, round (-angular) in outline. Perforations simple (Fig. 73). Intervessel pits opposite to alter­ nate, round and crowded, 6-20 11m in horizontal diameter (Fig. 72); without vestures. Vessel-ray pits with reduced borders, round (to horizontal or vertical), opposite, 4-15 11m in horizontal diameter (Fig. 73). Thick helical thickenings present throughout vessel elements (Fig. 73). Vessel element length 480-570 11m. Tyloses conspicuous (Fig. 72, 73). - Fibres with minutely bordered pits in radial walls, all septate (Fig. 72); walls thin to thick, 3.5 11m; 1140-1460 11m long. F/V ratio 2.1-2.5. -Axial paren­ chyma scanty paratracheal, 2-7 cells per strand, grading into septate fibres. - Rays heterocellular, 1-6 cells wide, up to 1.8 mm tall, rarely with occasional sheath cells (Fig. 72); perforated ray cells with simple perforations found in one specimen of T. tiliifolia (Suzuki & Noshiro 8540005); rays composed of procumbent, square, and upright body cells, and one row of marginal upright cells. - Crystals absent.

GARRYACEAE

Garrya - Fig. 56-58 Growth rings distinct (Fig. 56), marked by differences in vessel diameter between latewood and earlywood and radially flattened fibres at the end of growth rings; rare­ ly indistinct in G. fadyenii (SJRw 16712, 19632) and G. laurifolia (MADw 23809). Wood diffuse-porous, with a gradual decrease in vessel diameter towards the growth ring boundaries in temperate species (Fig. 56), with a tendency to semi-ring-porosity in G. buxifolia. - Vessels mostly solitary or rarely in multiples or clusters of 2 or 3, vessel grouping index 1.00-1.07, 43-362/mm2, 16-53 !lffi in tangential diameter, round-angu1ar in outline. Perforations scalariform with 3-8 bars (Fig. 58), more than 6 in the subtropical species G. fadyenii, G. laurifolia, and G. longifolia, of subg. Fa­ dyenia. Intervessel pits alternate-opposite, 3-6 !lffi in horizontal diameter; without vestures. Vessel-ray pits with distinct borders, occasionally unilaterally compound, round (to horizontal), opposite to alternate and crowded, 4-10 11m in horizontal diam­ eter. Distinct helical thickenings occur throughout vessel elements (Fig. 58). Vessel element length 400-830 11m, longer than 600!lffi in the three subtropical species and one G. elliptica specimen (MADw 45076). Purple gums occasionally present in in­ nermost rings. - Fibres with distinctly bordered pits in both radial and tangential walls, with distinct helical thickenings; walls thin to thick, 3-10 11m; 520-1670 11m long, longer than 1000 11m in the three subtropical species and one G. elliptica speci­ men. F/V ratio 1.2-2.1. - Axial parenchyma diffuse or diffuse-in-aggregates, and scanty paratracheal (Fig. 56), 4-8 cells per strand. - Rays heterocellular, 1-11 cells wide, up to 6.5 mm tall, composed of procumbent, square, and upright body cells,

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$M ~ III ~ ~ I m11l! ~ ~~ ~ ~ i-~ ~ ~ ~ ; ~ !t:: R' W:~ ~ Bi: }?ia i.

It ff' ~ h!l ~ ~R r\ t:il ~ - ~~ ~ IHI;t ~ ~r- ~ U ~ ~f1 MI[ ~ ~ _!=~ I~ ~. ~~ K 1 ~ ßI!"i ~ ~ ~ ~ Ir 62 ~ ~ IK 63 ____ :M..lll:oI--_ '" ~ K rl j!\~ ~ t< ~ rr- S

..; ~~ P'

~ ~ ~ ~ ~ ~

~ 1~: ~ 'B ~ )' :t J; ~ ~

~ "} ~ ll$I4 II-~ - Fli ~~ ~ 6S ~ •- Fig. 62-64. Davidia involucrata (TWTw 3194) - 62: TS, diffuse-porous wood with vessel multiples.- 63: TLS, heterocellular rays. - 64: RLS, scalariform perforation with many bars and dense vessel-ray pits. - Fig. 65-67. Nyssa sylvatica (65, 66: MADw 1081,67: MADw 3027) - 65 : TS, diffuse-porous wood with many vessel multiples. - 66: TLS, uniseriate rays and opposite to scalariform intervessel pits. - 67: RLS, scalariform perforation. - Sc ale bars = 250 J.IID in Fig. 62, 65; 100 J.Im in Fig. 63. 66; 50 J.Im in Fig. 64, 67.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 69 with 1-3 marginal rows ofupright cells, with occasional sheath cells (Fig. 57); perfo­ rated ray cells with simple or 1 or 2 barred perforations occasionally present in G. flavescens, G. fremontii, and G. veatchii of subg. Garrya. Crystals absent.

NYSSACEAE

Camptotheca - Fig. 59-61 Growth rings distinct (Fig. 59), marked by smaller and fewer vessels and reduced radial diameter of fibres at the end of growth rings. Wood diffuse-porous (Fig. 59).­ Vessels solitary or in radial multiples or clusters of 2-3(-5), vessel grouping index 1.05-1.18, 64-80/mm2, 48-60 /lm in tangential diameter, round to angular in out­ line. Perforations scalariform with 17-21 bars (Fig. 61). Intervessel pits round and crowded, opposite (to alternate or scalariform), 6-7(-23) /lm in horizontal diameter (Fig. 60); without vestures. Vessel-ray pits with distinct borders, round and dense, 4-7 /lm in diameter, opposite to alternate (Fig. 61). Helical thickenings usually pres­ ent in vessel element tails. Vessel element length 760-1330 /lm. - Fibres with dis­ tinctly bordered pits in both radial and tangential walls (Fig. 60); walls thin to thick, 4-8 /lm thick; 1280-2010 /lm long. F/V ratio 1.5-1.8. -Axial parenchyma diffuse (Fig. 59), 4-8 cells per strand. - Rays heterocellular, 1-3 cells wide, up to 0.8 mm tall, composed ofprocumbent body cells and 1-4(-9) marginal rows of square or up­ right cells (Fig. 60). - Prismatic crystals common in 4-16-chambered axial paren­ chyma strands.

Davidia - Fig. 62-64 Growth rings distinct (Fig. 62), marked by differences in vessel diameter between latewood and earlywood and radially flattened latewood fibres. Wood diffuse-porous (Fig. 62). Vessels solitary or in radial multiples or clusters of 2-3(-5), vessel group­ ing index 1.08-1.16, 69-100/ mm2, 44-60 /lm in tangential diameter, round-angular in outline. Perforations scalariform with 60-91 bars (Fig. 64). Intervessel pits op­ posite to scalariform, closely spaced, 6-40 /lffi in horizontal diameter; without vestures. Vessel-ray pits with distinct borders, round and crowded, 3-4(-12) /lm in horizontal diameter, opposite (to scalariform) (Fig. 64). Helical thickenings absent. Vessel ele­ ment length 1020-1590 /lffi. - Fibres with distinctly bordered pits in both radial and tangential walls (Fig. 63, 64); walls thin, 2.5 /lm thick; 1500-2210 /lffi long. F/V ratio 1.4-1.5. Axial parenchyma diffuse(-in-aggregates), 6-10 cells per strand. - Rays heterocellular, 1-3 cells wide, 0.8 mm tall, composed of procumbent body cells with 1-15 marginal rows of upright or square cells (Fig. 66). - Crystals absent.

Nyssa - Fig. 65 -70 Growth rings distinct in temperate species (Fig. 65), marked by differences in ves­ seI size in latewood and earlywood and radially flattened latewood fibres; indistinct and faintly marked by thick-walled fibres, to absent in N. javanica (Fig. 68). Wood diffuse-porous (Fig. 65, 68). Vessels solitary or in radial multiples or occasional clus­ ters of 2-5(-9), vessel grouping index 1.11-1.88; 26-150/mm2 in temperate to sub-

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Fig. 68-70. Nyssajavanica (BZFw 9999) - 68: TS, diffuse-porous wood with vessel multiples and indistinct growth rings. - 69: TLS, heterocellular rays and opposite intervessel pits. - 70: RLS, scalariform perforation and vessel-ray pits. -Fig. 71-73. Toricellia arguta (SJRw 21773) - 71 : TS, ring-porous wood with larger solitary vessels at growth ring boundaries. - 72: TLS, heterocellular rays and septate fibres. - 73: RLS, simple perforations, distinct helical thickenings, vessel-ray pits reduced borders. - Scale bars =250 iJm in Fig. 68, 71; 100 iJm in Fig. 69, 72; 50 iJm in Fig. 70, 73.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 71 tropical species (Fig. 65), 11-23/mm2 in N. javanica (Fig. 68); 36-66 11m in tangen­ tial diameter in the former, 77-109 11m in N. javanica; round-angular in outline. Per­ forations scalariform with 20-50 bars (Fig. 67, 70). Intervessel pits opposite to scalariform, crowded, 4-6(-35) /Jffi in horizontal diameter (Fig. 66, 69); without vestures. Vessel-ray pits with distinct borders, round (to horizontal) and dense, 3-8 (-20) 11m in diameter, opposite to alternate (Fig. 67, 70). Helical thickenings usually present in vessel element tails, but lacking in some specimens. Vessel element length 850-1340 11m in temperate to subtropical species, 1360-2080 11m in N. javanica. Tyloses present only in one N. sylvatica specimen (MADw 15811); reddish purple gums occur occasionally in N. biflora (Aw 28096, MADw 8431) and one specimen of N. javanica (FPAw 26001). - Fibres with distinctly bordered pits in both radial and tangential walls, but fewer in tangential walls; occasionally septate in two specimens of N. sinensis (TWTw 7468, MADw 1081) or with dark purple deposits in one sampIe of N. sylvatica (MADw 15811), all septate to nonseptate and rarely with deposits in N. javanica (Fig. 69); walls thin to thick, 2.5-10 11m; 1120-2130 11m long in temper­ ate to subtropical species, 1790-2900 11m long in N. javanica ; fibres rarely as wide and as thin-walled as vessels; indistinguishable from the latter in cross sections in single specimens of N. aquatica (MADw 8482) and N. biflora (Aw 28096). F IV ratio 1.1-2.1. -Axial parenchyma diffuse(-in-aggregates), and occasionally scanty para­ tracheal (Fig. 65,68),4-14 cells per strand. - Rays heterocellular, 1-3 cells wide in temperate to subtropical species (Fig. 66, 69), up to 6 cells wide in two sampies of N. javanica (TWTw 4454, 12529); composed of procumbent body cells with 1-6 marginal rows of weakly procumbent, upright or square cells. - Prismatic crystals usually in 4-1O-chambered strands ofaxial parenchyma in temperate to subtropical species, in non-chambered or 2-16(-34)-chambered strands in N. javanica.

Key to the woods of the Cornaceae alliance la. Perforations mostly or exclusively simple ...... 2 b. Perforations exclusively scalariform ...... 4 2a. Perforations simple and occasionally scalariform ...... Melanophylla b. Perforations exclusively simple...... 3 3a. Helical thickenings throughout vessel elements ...... Toricellia b. Helical thickenings absent .... Alangium sect. Alangium, Marlea, Rhytidandra 4a. Helical thickenings throughout vessel elements ...... 5 b. Helical thickenings absent or restricted to vessel element tails ...... 7 5a. Helical thickening absent in fibres, perforations with more than 10 bars ... 6 b. Helical thickenings in fibres, perforations with up to 10 bars...... Garrya 6a. Rays up to and over 6 cells wide, usually in two sizes ...... Aucuba b. Rays up to 4 cells wide ...... Corokia 7a. Uniseriate rays common among the multiseriate rays ...... 8 b. Uniseriate rays absent or very rare ...... Aralidium 8a. Helical thickenings in vessel element tails ...... 9 b. Helical thickenings absent ...... 11

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9a, Perforations with at most up to 25 bars, , , , , ...... Camptotheca b. Perforations with 20 to 60 bars...... 10 lOa. Vessel-ray pits bordered, crowded ...... Nyssa b. Vessel-ray pits with reduced borders ...... Mastixia lla. Perforations with up to 15 bars...... 12 b. Perforations with more than 15 bars...... 13 12a. Apotracheal parenchyma in irregular lines, fibres with almost c10sed lumina .. · ...... Alangium sect. Constigma b. Apotracheal parenchyma diffuse-in-aggregates, fibres less thick-walled ...... · ...... Griselinia 13a. Apotracheal parenchyma diffuse or diffuse-in-aggregates, fibres non-septate 14 b. Apotracheal parenchyma absent or rare, fibres septate ...... Helwingia 14a Vessel-ray pits bordered, crowded, round ...... 15 b. Vessel-ray pits sparse, at least partly with reduced borders, round to horizontal · ...... Diplopanax 15a. Rays more than 3 cells wide ...... 16 b. Rays up to 3 cells wide ...... Davidia 16a. Crystals usually absent ...... Cornus b. Prismatic crystals common in ray cells ...... Curtisia

Latitudinal trends and correlation among quantitative characters Latitudinal trends in the wood structure of Cornales are c1ear in large genera such as Cornus, Alangium, and Garrya. When there was limited information ofthe locality associated with the wood sampies, a latitude was assigned based on information on the species in various floras or floristic databases. Five quantitative characters, i.e., vessel element length, fibre length, vessel frequency, tangential vessel diameter, and number of bars per perforation, were plotted against latitude. Data of tree size were very meagre and were not directly comparable with quantitative characters. In Cornus and Alangium, tree size was categorised directly from collection records or deduced from floristic literature; plants were divided into shrubs (multistemmed, height up to 3 m), small trees (3-10 m), and large trees (over 10 m tall). These size c1asses were compared with quantitative anatomical data. Because the real tree size was unknown, only simple correlations with wood anatomical characters are studied, and multiple regression together with latitude has not been attempted. The latitudinal trends have so far been interpreted in relation to macroclimate or vulnerability. This will be discussed with an analysis of trends in East Asian Cornus species in a separate paper (Noshiro & Baas, in preparation). Here latitudinal trends are studied to assess variation in quantitative characters within the taxa and to explore whether different character states can be meaningfully defined for c1adistic analysis in these continuously varying features. Cornus has a comparatively homogeneous wood structure and shows c1ear lati­ tudinal trends in several characters. Localities of Cornus specimens range from the equatorial tropics to 54 0 northern latitude; only two of the species studied, C. volkensii and C. peruviana, occur in the equatorial region of the southern hemisphere. Vessel element length and fibre length both have a negative linear correlation with latitude

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(Fig. 74). In both characters, subg. Syncarpea tends to have higher values than the regression lines, but the other sub genera closely follow these lines. Vessel element length and fibre length respectively decrease from ab out 1600 J..lm and 2400 J..lm near the equator to half those lengths at around 50° latitude. In relation to tree size, large tree species tend to grow at lower latitudes and have longer vessel elements, whereas shrubby species tend to grow at higher latitudes and have shorter vessel elements (Fig. 75). However, in each size class there is a negative correlation with latitude. Nearly the same negative linear correlation exists between tangential diameter of vessels and latitude: tangential diameter of about 80 11m around the equator halves at around 50° latitude (Fig. 74). Vessel frequency shows a more or less exponential in­ crease against latitude, but the trend above 40° is biased upward with small dense vessels often associated with shrubby species. Vessel frequency seems to be corre­ lated more with tree size than with latitude: larger trees clearly have fewer vessels, and latitudinal trends within each size class are not so clear (Fig. 75). Bar numbers tend to have characteristic ranges for the sub genera irrespective of latitude; subg. Afrocrania, Mesomora, and Syncarpea mostly have values above 40, and subg. Kra­ niopsis and others have values below 40. There are, however, some exceptions in subg. Kraniopsis (Fig. 74). In Alangium, two distinct groups were recognised. Provenances of Alangium speci­ mens range from 28° southem latitude to 36.5° in the northem hemisphere. Section Constigma, which is distinct in this genus in having scalariform perforations, has distinctly the longer vessel elements and fibres, l300-2000 and 2500-3200 J..lffi re­ spectively (Fig. 76). Species of sect. Constigma are large trees usually over 20 m tall and are restricted to low latitudes, but the other sections also include trees of similar size such as A. griffithii, A. longiflorum and A. kurzii, and the anatomical differences are thus significant. Sections Marlea, Alangium, and Rhytidandra occupy a common range in both vessel element length and fibre length, and these ranges have a weak negative correlation with latitude. The rate of reduction in length is far less than in Cornus, and temperate species have vessel elements and fibres that are about 20-30% shorter than those of the tropical species. Vessel frequency increases with latitude as in Cornus, but the trend is obscured with scattered values of sect. Marlea, Alangium, and Rhytidandra (Fig. 76). This feature is significantly correlated with tree size: large trees have vessels less than 20/mm2, and shrubs or small trees have more numerous vessels (Fig. 77). Tangential vessel diameter does not have any correlation with lati­ tude, and sect. Marlea shows a wide variation covering the whole range. InA. chinense, it ranges from 75-180 11m between 20-30° and 70-140 11m between 0-10°, and the latitudinal trend is non-existent. Vessel diameter shows an overall correlation with tree size. Trees with semi-ring or ring-porous wood tend to have exceptionally wide vessels (Fig. 77). Vessel element lengths of Alangium sect. Constigma overlap with the upper ranges of Cornus, and ranges of Nyssaceae overlap entirely with Cornus; jointly they show a rather steep decline with increasing latitude (Fig. 78). Values of the other sections of Alangium, and of Garrya clearly come lower than these main groups, and decline (text continued on page 80)

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Cornus "subg. Afrocrania & Mesomora (/mrn~ o subg. Kraniopsis 240 " subg. Syncarpea • others G-200 ~ f(x) = -16.7 x + 1644 ~ 160 1800 0"' R= -0.721** cf' ~ 120 " " ~ ...... ,..s::::: 1600 • ". Cl) ~ Ul 80 00 o Ul o .. Cl) ~ o .... 1400 • > 40 j 0" ~ -"'" . . 0 ~ ...... • 0 0 10 20 30 40 50 60 j 1200 0· ...... r!' • (11m) '.,0.11 ~ • a 'cP o 00 0..... • .... 120 -0.73 76.9 Q) 1000 o o o.·~ .. o ...., ...... choo(tj ~ 800 0 00

600+-~r-~~-.~-r~.-~ o 10 20 30 40 50 60

{j.un) 2400 f(x) = -25.9 x + 2448 • R= -0.715** 2200 o .. 10 20 30 40 50 60 ."'0 2000 .. • 0 • ci'o. 80 \~ 'to 18OO ~" '.. ie 0 • 70 ~ 0,. • j 1600 o ~~'. ~. 0 •• ~ 60 ..0 " Cl) °

800 10+-~~-.~.-~~-.~~ 0 10 20 30 40 50 60 o 10 20 30 40 50 60 Latitude Latitude

Fig. 74. Vessel element length, fibre length, vessel frequency, tangential diameter, and bar number of Cornus against latitude. Each dot shows the mean value for a single specimen. Significance level: ** = 0.5%, * = 1%.

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Cornus Vessel element length (/.lrn)

2000 2000 1500 1500 1000 1000 500 500 0

Shrub

Latitude Small-Large Tree 60 Large Tree

Vessel frequency (/ rnrn 2)

240 240 200 200 160 160 120 120 80 80 40 0 Shrub

Small Tree Latitude 60 Small-Large Tree Large Tree Fig. 75. Vessel element length and vessel frequency of Cornus against latitude and tree size.

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Alangium • sect. Constigma o sect. Marlea • (A. chinense) * sect. Alangium & Rhytidandra

(1J.l11)

2500 o

>.. 2000 .... u 40 .. ~ 0 ...... Q) i .. .. ;j 0 "E 1500 .... 0'" 30 Q) f(x) = -7.28 x + 748 Q) • • I-< • * S R = -0.526" ~ Q) ...... • 8 ...... 1000 Q) 20 .. ~ CI) .. CI) • ...... o 0 Q) Q) ...... • • CI) .. * • oi CI) 500 > 10 • J i Q) .. , • 0 o • .. •• • > 0 0 0 0 10 20 30 40 0 10 20 30 40 Latitude Latitude

(~m) (~m) 3500 180 • .. I-< .. ...Q) 160 • 3000 ...... Q) .. .. S ...... ctl 140 • ...... • 0 ..r::... 2500 a .... ~ 0 • ...... ~ Q) 120 Q) CI) o..' • ., ~ .. • 0 CI) ....J 2000 Q) ~~ Q) • • ~ * • , .. 0 I-< ..•..•Qj, . •• 0 > 100 • ••.. "Ir.. • 0 ...... t ..0 o 0 ctl ...... • oil 0 1500 • 0 .'i ...... • ~ * o • ... • * ~ 80 .. 0 • 0 Q) • 0 • • • 0 * 0 bJ) • •• 0 1000 0 ~ f(x) = -9.68 x + 1733 ctl 60 R = -0.412' E-< • • 0 500 40 0 10 20 30 40 0 10 20 30 40 Latitude Latitude

Fig. 76. Vessel element length, fibre length, vessel frequency, and tangential diameter of Alan­ gium against latitude. Each dot shows the mean value for a single specimen. Significance level: ** = 0.5%, * = 1%.

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Alangium Vessel frequency (/mm2)

50 50 40 40 30 30 20 20 10 10 o Shrub

Small Tree Latitude Small-Large Tree 40 Large Tree

• diffuse-porous .. semi-ring-porous & ring-porous Tangential vessel diameter (!lm)

200 200 160 160 120 120 80 80 40 40 o o o Shrub

Small Tree

Latitude Small-Large Tree 40 Large Tree

Fig. 77. Vessel frequency and tangential diameter of Alangium against latitude and tree size.

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()Jm) • Alangium (/nun2j 2800 .. Cornus 400 o Mastixia " Garrya * Nyssaceae 350 ..t:: 2400 + others • o cA> ..... o >-,300 ~ u • 0 j 2000 • 0 00 + s:: o~ Q) 250 + + •• 0 ;J + ~ cr' • + • • 2 - • + alt. 1600 I • ~ 200 j (.,I.., • ... ·\, ...... ~ I -&l 150 + • CIJ 'i 1200 Q) CIJ . + :> 100 ~ . • 800 · ..• . • • a. • 50 I'· 20 30 40 50 60 10 20 30 40 50 60

()Jm)

4000 160

140 3500 o ~ .... o 80 + • 0 .: 0 oE 3000 .",!> ~ · .­ -&l100 ;·0· . + CIJ j 2500 Q) + Q) •+ .. 80 I-< - > tt: 2000 ...... 60 1500 · · . 40 1000 20

O+-~~-.~.-~~-.-r, 20 30 40 50 60 o 10 20 30 40 50 60 Latitude Latitude

Fig. 78. Vessel element length, fibre length, vessel frequency, and tangential diameter of all the genera against latitude. Each dot shows the mean value for a single specimen.

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90 * 80 • Alangium .. Cornus 70 + 0 Mastixia ~ Q) 0 • Garrya 60 00 0 Nyssaceae 0 0 • * + * others "S 0':9 + + ;:::l 50 8 ... ~ ...." g'lo 0 " Z *00 0 .... " 11 * ... + ... ~ 0 ... t " ce 40 .' 0 + ...... * "'* 4iIf . (:0 " *:.\ ~'*t" " " 30 • .' It + + .t r. ~ ... :.t.!... JI. 20 1**.... \ ...... + * +... ++ •• + "+ 10 : : . • + + • ttI • • 0 ...... "' .... 0 10 20 30 40 50 60 Latitude (J.1ffi) 180 • Nyssaceae f(x) = 2875000 x-2.66 R2 = 0.814 160 • , :* Mastixia f(x) = 324.0 x-0370 R2 = 0.402 140 *: other taxa f(x) = 323.4 x.()·461 "t 120 R2 = 0.854

Q) 100 CIJ CIJ Q) > 80

". + + 20 *:++ + + •

O~----.-----.-----.-----.-----.-----.-----.----. o 50 100 150 200 250 300 350 Vessel Frequency

Fig. 79. Bar number against latitude and tangential diameter against vessel frequency for all the genera.

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()lIIl) • Alangium more gradually with increasing '" Cornus latitude. Values of Mastixia oc­ 3000 0 Mastixia • Garrya cupy a distinct range above * Nyssaceae 0 these groups, and do not show + others 0 a clear latitudinal trend. In fibre 2500 00 length, the upper range of Cor­ 00 't9 0 0 0 nus comes into the gap between ...c:..... 0 .0 sections of Alangium, and its 0 gp 2000 + 0 0 Q) 0 middle range overlaps with the .....l •• 0 0 .. lower range of Alangium ex­ ..... o • ~ + .t Q) 0 •• cluding sect. Constigma. Values +11. * JJ.. • • S 1500 ... .. of Garrya mostly come lower •• 0 .' • Q) ~ ... + Mr,... •• p;.j • • than those of Cornus, and show - .. ~ ...+ .. +++ Q) ... 111... + + a sirnilar decline as Cornus with -rJl rJl ~.t.!_. Q) 1000 .~ * • increasing latitude. The range of 1+ .. ..." -. > .,ll. itY·'. Mastixia mostly comes above +:}.,. tlo<· - • • \ .... ." • that of the other taxa. Vessel fre­ +11.-- -- ,1· •. • • •• 500 .,. + • • • •• • • quency seems to show an expo­ - . nential increase with latitude, but is strongly influenced by the

O~~~~-.~~-r~~,-~ high frequencies in subtropi­ o w ~ ~ W 100lW1~1~ cal to temperate shrubby taxa, Tangential Vessel Diameter ()lm) especially Garrya, and the real, intrinsic latitudinal trend is not Fig. 80. Vessel element length against tangential diam­ clear. Though strongly disturb­ eter for all the genera. ed by the wide ranges of Alan­ giuminrniddlelatitudes,tangen­ tial vessel diameter seems to have a negative correlation with latitude along the ranges of Cornus, Mastixia, and Nyssaceae. Values of Garrya and most shrubby genera, i. e., Aucuba, Corokia, Griselinia (excluding G. lucida), and Helwingia, are below this main trend, and are associated with their shrubby habit. Bar numbers do not show any clear latitudinal trend within the respective taxa, and the taxa can be divided into those with over 20 bars and those with less than 20 bars (Fig. 79). Vessel frequency and vessel diameter of a11 the studied taxa are mutua11y strongly and negatively correlated: the relation fo11ows an exponential curve (Fig. 79). This trend is largely caused by Alangium, Cornus and Garrya in order of increasing vessel frequency. Mastixia and Nyssaceae have a clearly different relationship of these values: Mastixia shows a more gentle, less significant trend, and Nyssaceae shows a steeper exponential curve. Tangential vessel diameter also has a linear correlation with vessel element length, but Alangium has distinct trends quite different from the other taxa (Fig. 80). In the two groups within this genus, vessel element length and fibre length both show a gentIer increase against tangential vessel diameter compared with the main group consisting largely of Garrya, Cornus, and Nyssaceae. Vessel ele­ ment length and fibre length of Mastixia have a slightly steeper increase with vessel diameter than the other genera.

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PHYLOGENETIC ANALYSIS

A phylogenetic analysis was carried out based on a datamatrix of 21 selected charac­ ters that showed discrete variation among the taxa studied (Table 3). Characters di­ rectly influenced by habitat such as distinctness of growth rings and tyloses, and those without clear distinction in character states in the Comales such as porosity, two ray sizes or sheath cells in rays were not employed. Of dependent characters such as bordered pits in both radial and tangential walls of fibres versus simple to rninutely bordered pits only in radial walls, only one was selected (character 10). Autapomorphic characters such as marginal parenchyma and druses in sect. Marlea of Alangium or absence of uniseriate rays in Aralidium were excluded. Quantitative characters, i.e., the vessel grouping index (character 1), vessel fre­ quency (character 2), tangential diameter ofvessels (character 3), number ofbars per scalariform perforation (character 4), vessel element length (character 8), and F/V ratio (character 13), were divided into character states considering the distribution of mean values of all specimens for genera or infrageneric taxa. Large genera, such as Comus, Alangium, and Garrya, had specific and continuous ranges of variation against latitude, and the different dependencies must also be phylogenetically informative. The division into character states may, however, be artificial. Thus coding was based on the gaps detected between groups of measurements. In vessel grouping index (char­ acter 1), the division within Comus is considered as a criterion for recognising char­ acter states. Vessel frequency (character 2) has two main gaps at c. 40 and 100/mm2 (Fig. 78), but the lower one is traversed by the range of Comus, and only the upper one was adopted. Though strongly correlated with vessel frequency, tangential diam­ eter of vessels (character 3) has a gap at 40 Iilll that corresponds with the lower limit of Alangium and Comus and the upper one of Garrya (Fig. 79). In bar number (char­ acter 4), the upper division within Comus coincides with ranges of Mastixia and Aucuba, and the lower division includes few-barred taxa such as Alangium sect. Constigma, Corokia, Griselinia, Garrya, and Camptotheca (Fig. 79). In Comus, ves­ seI element length (character 8) and fibre length showed a strong linear correlation with latitude without any disjunction or sudden changes, and the distinction between subgenera was difficult (Fig. 74). In Alangium, both ofthese characters were distinct­ ly longer in sect. Constigma than in other sections. The gap in vessel element length between the sections of Alangium nearly conformed to the range of Comus, and the range of fibre length in the temperate species of Alangium nearly overlapped the mid­ dIe range of Comus (Fig. 78). Thus, in vessel element length, character states are se­ lected not to divide the continuous ranges of Comus, but to distinguish two separate ranges of Alangium, one above that of Comus and another below it. Fibre length was excluded from the datamatrix because of the strong correlation with vessel element length. The F/V ratio (character 13) was below 2.0 and indistinguishable for almost all taxa except for three sections of Alangium and Toricellia that have two to four times longer fibres than vessel elements and is divided into states at this value. Qualitative characters were divided into character states on the one hand, largely following the IAWA list (1989), and on the other to exclude autapomorphy in the total

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Table 3. Datarnatrix for the cladistic analysis.

character Taxon 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Alangium -Alangium 1 0 1 3 1 1 0 2 0 1 0 0 I 2 0 I I I 0 I I Alangium- Constigma I 0 1 2 I 1 0 0 0 I 0 0 0 2 0 I I I 0 I 1 Alangium-Marlea 1 0 I 3 I I 0 2 0 I 0 0 I 2 I I I I 0 I I Alangium -Rhytidandra I 0 1 3 1 1 0 I 0 1 0 0 0 2 0 1 I I 0 1 0 Aralidium I 0 1 I 0 1 0 1 0 I 0 1 0 0 1 0 2 0 0 0 0 Aucuba 0 0 0 Oll 0 0 2 I I I 0 1 0 0 I 2 0 0 0 0 Cornus-Afrocrania 1 0 I 0 0 0 0 I 0 0 0 0 0 2 0 I I 0 0 0 Cornus-Cornus 0 0 1 1 0 0 0 I 0 0 0 0 0 2 0 I I 0 0 0 Cornus- Cynoxylon 0 0 1 1 0 0 0 I 0 0 0 0 0 2 0 I I 0 0 0 Cornus-Discocrania 0 0 I I 0 0 0 1 0 0 0 0 0 2 0 1 I 0 0 0 Cornus-Kraniopsis 0 0 I 1 0 0 0 I 0 0 0 0 0 2 0 I I 0 0 0 Cornus-Mesomora I 0 1 Oll 0 0 0 1 0 0 0 0 0 2 0 1 I 0 0 0 Cornus-Syncarpea 0 0 1 0 0 0 0 1 0 0 0 0 0 2 0 1 1 0 0 0 Cornus-Yinquania 0 0 I 1 0 0 0 I 0 0 0 0 0 2 0 1 I 0 0 0 Corokia 0 1 0 I 0 0 2 2 I 1 0 1 0 0 0 ? I 0 0 0 0 Curtisia 0 0 I I 0 0 0 0 0 0 0 0 0 2 0 1 1 I 0 I 0 Diplopanax 0 1 1 1 0 1 0 I 0 0 0 0 0 2 0 I I I 0 0 0 Griselinia o 0/10/1 2 0 1 0 1 0 0 0 1 0 2 0 0 2 0 0 1 0 Helwingia I I 0 I 0 I 0 1 0 I 0 1 0 0 1 0 2 0 0 0 0 Mastixia-Manglesia 0 0 1 0 0 I I 0 0 0 0 0 0 2 I I I 1 0 0 0 Mastixia-M.-Alternae 0 0 I 0 0 I I 0 0 0 0 0 0 2 I I I I 1 0 0 Mastixia- M. -Oppositae 0 0 I 0 0 I I 0 0 0 0 0 0 2 I I I I 1 0 0 Melanophylla 1 0 I 3 0 I 0 I 0 I 0 I 0 0 I 0 I 0 0 0 0 Toricellia I 0 1 3 0 I 2 2 0 1 0 I I 0 I 0 I 0 0 0 0 Garrya -Fadyenia o 0/10/1 2 1 0 2 2 0 0 1 0 0 2 I 0 2 0 0 0 0 Garrya -Garrya 0 I 0 2 1 0 2 2 0 0 1 0 0 2 1 0 2 0 0 0 0 Camptotheca I 0 1 I 0 0 1 I 0 0 0 0 0 1 0 I 0 I 0 0 I Davidia I 0 I 0 0 0 0 1 0 0 0 0 0 I 0 1 0 I 0 0 0 Nyssa spp. 1 0 1 1 0 0 1 1 0 0 0 0 0 1 0 1 0 1 0 0 I Nyssa javanica 1 0 I 0 0 0 I 0 0 0 0 1 0 1 0 1 Oll 1 0 0 1 Hydrangea paniculata 0 0 I 0 0 0 0 0 0 0 0 I I 0 0 1 0 0 0 Hydrangea shrub 0 I 0 0 0 0 0 I 0 1 0 0 0 ? 1 0 0 0 0 Cercidiphyllm 0 1 I 0 I 0 0 0 0 0 0 0 0 0 0 0 Daphniphyllm 0 1 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 Hamamelis 0 0 2 0 0 0 0 0 0 0 I 0 0 0 0 0 0 1. Vessel grouping index: 0=< 1.1; 1 = :2: 1.1 2. Vessel frequency: 0=< 100/mm2; 1 = :2: 100/mm2 3. Tangential vessel diameter: o = < 40 11m; I = :2: 40 11m 4. Perforation (number of bars): 0= :2: 40; I = 15- < 40; 2 = < 15; 3 = simple/mixed 5. Intervessel pits: o = opposile-scalariform; 1 = alternate 6. Vessel-ray pits: o = bordered; 1 = with reduced borders I apparently simple 7. Helical thickenings: 0= absent; I = in vessel element tails; 2 = throughout body ofvessel 8. Vessel element length: 0=:2: 1600 11m; 1 = 800- < 1600 11m; 2 = < 800 11m [element 9. Vascular tracheids: o = absent; I = present 10. Fibre pils: o = distinctly bordered; 1 = simple to minutely bordered 1/. Helical thickenings in fibres: o = absent; I = present 12. Septate fibres: o = absent; 1 = present 13. F IV ratio: o = < 2.0; 1 = :2: 2.0 14. Apotracheal parenchyma: 0= absent/rare; I = diffuse; 2 = diffuse-in-aggregates 15. Paratracheal parenchyma: 0= absent/rare; 1 = scanty/vasicentric 16. Maximum strand length: o = ::; 8 cells; 1 = :2: 9 cells 17. Ray width: o = ::; 3 cells; I = 4-9 cells; 2 = :2: 10 cells 18. Ray body: 0= includes square/upright cells; 1= consists exclusively of 19. Axial canals: o = absent; I = present [procumbent cells 20. Prismatic crystals in rays: 0= absent/rare; 1 = common 21. Prismatic crystals in axial parenchyma: 0 = absent/rare; 1 = common

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 83 datamatrix. For instance, apotracheal parenchyma (character 14) in Alangium ranged from diffuse to in irregular lines depending on the species (Table 2). 'Diffuse-in-ag· gregates' was selected for this character for all the sections because this state is basic in most species and plesiomorphic for all sections and because 'in irregular lines' found only in tropical species results from well-developed diffuse-in-aggregates pa­ renchyma. Vasicentric to confluent parenchyma occurs only in Alangium sect. Marlea and is autapomorphic within Cornales. 'Scanty paratracheal' was therefore allotted to this section considering its development into 'vasicentric' as a general trend in wood evolution. Presence or absence of septate fibres (character 12) and distribution of para­ tracheal parenchyma (character 15) showed two equally frequent states in Nyssa spp. and N. javanica, and plesiomorphic conditions deduced from preliminary eladistic analyses are allotted to these characters in Nyssa. Presence of crystals (characters 20 & 21) was constant for most specimens within a taxon, and its absence may result from physiological conditions of specific individuals or the sampling point within a shrub or tree. Thus the representative state for the majority of specimens is used for that taxon. Ray composition (character 18) did not fit the division of the IAWA list, and the bodies of multiseriate rays sporadically ineluded square or upright cells in several taxa, but not throughout the ray (feature 109 of the IAWA list). Cell composition of multiseriate ray bodies shows continuous variation from Heterogeneous Type I to Heterogeneous Type HA of Kribs (1935) and was divided into ray bodies consisting exelusively of procumbent cells and those ineluding square or upright cells beside procumbent ones. Sheath cells have two extremes in Cornales, but continuously vary from one character state to another. Thus this character is exeluded from the datamatrix because division into distinct states could only be artificial. Cercidiphyllum, Daphniphyllum, and Hamamelis were chosen as outgroups. There is no consensus on the si ster group of the Cornales, but analysis of rbcL sequence data suggested taxa elose to the basallineage of Rosidae ineluding some Hamamelidae, i.e., Cercidiphyllum, Daphniphyllum, and Hamamelis (Chase et al. 1993). Among the genera used as an outgroup in the analyses of rbcL sequence data (Xiang et al. 1993), Cercidiphyllum, Daphniphyllum, and Hamamelis produce ample secondary wood that is comparable with the completely woody genera of Cornales, whereas the others are either incomparable herbs or shrubs with specialised woods. As representatives of Hydrangeaceae, which are considered elose to the Cornaceous elade (Xiang et al. 1993; Xiang & Soltis in press), we llsed Hydrangea serrata (Thunb. ex Murray) Sero and H. yayeyamensis Koidz., two shrubby species, and H. paniculata Sieb. & Zucc., a small tree. Heuristic search with equal weight for all characters produced twelve equally par­ simonious trees (Fig. 81, 82). All characters were unordered and the ACCTRAN op­ tion was used. Heuristic search was performed with random addition sequence, TBR branch-swapping and MULPARS option. Tree length was 70 steps, and the consist­ ency and retention indices were 0.386 and 0.744, respectively. Reweighting with rescaled consistency indices resulted in the same twelve trees. Ca1culation ofbootstrap percentage with equal weight for all the characters was beyond the ability of the

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Camptotheca 76 Nyssa spp. 64 Nyssa javanica ,1L ~ Davidia Cornus-Afrocrania ,L Cornus-Mesomora Cornus-Syncarpea Cornus- Cornus ,1JL Cornus-Cynoxylon Cornus-Discocrania I Cornus-Kraniopsis Cornus-Yinquania -.1L Alangium-Alangium 64 Alangium-Marlea 91 Alangium-Constigma 42 Alangium -Rhytidandra 2L JL ~ Curtisia Mastixia-Manglesia 63 Mastixia-M. -Alternae ~ Mastixia-M.-Oppositae Diplopanax Aralidium 56 Melanophylla ~ Toricellia ,lL Aucuba Corokia - ,-iL Helwingia 11 Hydrangea shrub --.lL Griselinia 47 Garrya-Fadyenia ~ Garrya-Garrya Hamamelis Hydrangea paniculata Daphniphyllum Cercidiphyllum

Fig. 81. Strict consensus tree of the twelve most parsimonious cladograms resulting from heuristic search.ltalic numbers indicate the percentage occurrence of each monophyletic group in the results of 100 bootstrap sampies.

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 85 c1adistic software, and no feasible results could be obtained. Thus boots trap percent­ ages for 100 replicates were calculated by choosing characters with equal prob ability, but reweighting them with the maximum value of rescaled consistency indices. In all the twelve trees obtained with equal weight, Cercidiphyllum comes at the base, and Daphniphyllum and Hydrangea paniculata at the next basal branches (Fig. 81). All trees support monophyly of two c1ades referred as c1ades land 11: c1ade I consisting of Alangium, Cornus, Curtisia, Diplopanax, Mastixia, and Nyssaceae, and c1ade 11 of all the other genera of Comales, Hydrangea shrubs, and Hamamelis. These two c1ades form a polytomy with Daphniphyllum and Hydrangea paniculata. The twelve trees differ in the branching above Griselinia within c1ade 11 and the basal positioning of Daphniphyllum and Hydrangea painiculata. In c1ade I, Diplopanax forms the basal branching to the two c1ades, one conisisting of Cornus and Nyssaceae and the other of Alangium, Curtisia, and Mastixia. Nyssaceae is paraphyletic and nested in sections of Cornus. In c1ade 11 Hamamelis forms the basal branching to a monophyletic c1ade consisting of all the other Comaceae allies and Hydragea shrubs. Clade I, consisting of Alangium, Cornus, Curtisia, Diplopanax, Mastixia, and Nyssa­ ceae, is defined on an apomorphy, parenchyma strands over 9 cells long (character 16), and a parallelism, ray width between 4-9 cells (character 17), and is weakly sup­ ported by a bootstrap percentage of 33% (Fig. 81, 82). Clade 11 is based on an apo­ morphy, tangential vessel diameter less than 40 f11ll (character 3), and a parallelism, perforation plates with less than 15 bars (character 4) with a bootstrap support of 47%. The monophyletic c1ade consisting of Comaceae allies and Hydrangea shrubs in c1ade 11 is based on two apomorphies: ray width over 10 cells (character 17) and ray body inc1uding square or upright cells (character 18) with a bootstrap support of 71%.

DISCUSSION

Character transformation in Cornales and systematic implications Wood anatomical features often show parallelisms and reversals and fail to resolve phylogeny. In Oleaceae, Baas et al. (1988) found parallelisms and revers als in all the 15 characters used for a c1adistic analysis. The 15 c1adograms they obtained inc1uded several polytomies and the resolution was very p0of. However, their c1adograms supported the monophyly of the subfamily Oleoideae, and were in c10se agreement with cytological and phytochemical information. In Rosaceae, Zhang (1992) found 14 parallelisms and/or reversals in Rosideae and Spiraeoideae p. p. and 16 in Quil­ lajeae and Prunoideae among the 18 characters used for a c1adistic analysis. His first analysis, covering the whole farnily Rosaceae, ended in an overflow of c1adograms, and he divided the datamatrix into three parts which were connected after indepen­ dent analysis. In this analysis, the small number of taxa and the critical assessment of character states seems to lead to a rather conc1usive result with a limited number of equally parsimonious trees. However, the consistency index is low due to frequent parallelisms and several reversals, as is common in wood anatomical features (Baas & Wheeler

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Camptotheca Nyssa spp. Nyssa javanica Davidia Comus-Afrocran ia Comus-Mesomora 1.0 Comus-Syncarpea Comus-Comus 6 Comus-Cynoxylon 1.0 Com us- Discocran ia I Comus- Kraniopsis 2 Comus- Yinquania Alangium-Alangium 15 Alangium-Marlea 0.1 Alangium-Constigma Alangium-Rhytidandra

0»1 0",1 Curtisia 1.0 4 715 Mastixia-Manglesia

1»0 0»10",1 Mastixia-M.-AItemae Mastixia-M.-Oppositae Diplopanax Aralidium Melanophylla 7 8 13 Toricellia 41014 0»21»2 0»1 Helwingia 2»10»1 2»0 2 15 12 6 7 9 Aucuba 1»00»1 8 17 Ir 1»00»20»1 Corokia 417 1»22»1 Hydrangen shrub 20 1»02,,1

0»1 Griselinia 5 6 7 81115 Garrya-Fadyenia »11»00»21»20»1 0»1 Garrya-Garrya

21415 Hamamelis Hydrangen paniculata 1»02»1 0»1 Daphniphyllum Cercidiphyllum

I apomorphy " parallelism X revers al

Fig. 82. One of the twelve most parsimonious cladograms resulting from heuristic search. Changes in character states are shown with character numbers in bold letters and character states below them. For characters and their states, see Table 3.

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1996). There were hundreds of poorly resolved trees one step longer than the mini­ mum-length trees. Thus the c1adogram based on wood anatomical features is far from robust. Among the 21 characters used for the c1adistic analysis of Comales, homoplasy was absent in five characters: presence of vascular tracheids (character 9), helical thickenings in fibres (character 11), maximum strand length (character 16), cellular composition of the rays (character 18), and presence ofaxial canals (character 19) (Fig. 82). Two other characters, vessel diameter (characters 3) and ray width (charac­ ter 17) also contribute greatly in defining c1ades land 11. However, smaller tangential diameter of vessels are common characteristics in shrubs compared with trees (Carlquist & Hoekman 1985; Fahn et al. 1986; Zhang et al. 1992) and contribute to the segre­ gation of shrubby taxa in this analysis. Even in c1ade 11, taxa of larger habit, i.e., Aralidium, Melanophylla, and Toricellia, have areversal to larger vessels, and this also indicates a correlation of vessel size with habit. Wide rays and ray composition (characters 17 & 18) characterise c1ade 11 along with the division into two distinct ray sizes or absence of uniseriate rays occurring in the terminal taxa, mostly as autapomor­ phy. Though the ray cell composition found in c1ade 11 is considered primitive in the phylogenetic schemes of ray types (Kribs 1935; Carlquist 1988), this seems to be a specialisation in this c1ade derived from rays with bodies consisting of procumbent cells and 1-6 rows of upright or square marginal cells, judging from the ray type of the outgroups. There are some reversals in key features that contradict the general trends in tracheary element evolution of dicotyledons established by Bailey and Tupper (1918) and Frost (1930). In the c1adogram, vessel element length (character 8) shows two parallel increases to over 1600 JJm and one parallel decreases to less than 800 JJffi in c1ade land three parallel decreases in c1ade 11 (Fig. 81). Vessel diameter (character 3) decreases to less than 40 JJffi in c1ade 11 once and increases once as areversal. Bar number per perforation (character 4) shows several reversals. Perforation plates be­ come simple in Alangium and again recover scalariform bars in Alangium sect. Con­ stigma, and bar number becomes less than 40 once and recovers to over 40 in c1ade 11 (Fig. 82). Fibre pits (character 10) become reduced to apparently simple in Alangium and once more in c1ade 11. Several reversals against the Baileyan trends c1early occur in tropical taxa in c1ade I, such as Alangium sect. Constigma, Nyssa javanica, and Mastixia, and may imply correlation with c1imatic or latitudinal circumstances or with their habit as large trees. However, this correlation is not c1ear in c1ade 11, and reversals occur sporadically in many c1ades. In this c1ade, reversals are correlated more with habit as discussed above.

A preferred delimitation of Cornaceae and related families The delimitation of Comaceae inferred from the wood anatomy inc1udes Cornus, Curtisia, Diplopanax, and Mastixia of Comaceae, Camptotheca, Davidia, and Nyssa of Nyssaceae, and Alangium of Alangiaceae. The other genera should be exc1uded from Comaceae.

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The above delimitation of Comaceae agrees with that of Eyde (1988) except for his exclusion of Alangium and Curtisia from Comaceae (Eyde & Xiang 1990). He treated Cornus and Mastixia as monophyletic and placed Nyssaceae as their si ster taxon. He was not sure about the inclusion of Alangium in Comaceae and discussed its putative position in his Comaceous diagram, sister to Cornus, or to Cornus & Mastixia. He seemed ready to accept its inclusion in Comaceae with additional evi­ dence. Eyde (1988) definitely excluded Curtisia from Comaceae due to its inferior ovary and central bundles in its ovary against superior ovary and transept bundles in Comaceae. Mainly because of similarities in fruit structure, Eyde & Xiang (1990) placed Diplopanax in Mastixioideae together with Mastixia and considered Cornus to be their si ster group, based on two-armed hairs. The delimitation ofComaceae based on wood anatomy mostly conforms to that deduced from rbcL sequence analyses in the inclusion of all the eight genera in clade I (Xiang et al. 1993; Xiang & Soltis in press). They recognised four lineages: the Cornus-Alangium group, Curtisia, the Nyssoid-Mastixioid group (Nyssa, Camptotheca, Davidia, Diplopanax, Mastixia), and the Hydrangeaceae-Loasaceae group in the Comaceous clade, but the position­ ing of these four groups was not resolved. The Comaceous clade based on rbcL se­ quence analyses differs from the present delimitation of Comaceae in the monophyly of the Nyssoid-Mastixioid group and that of Cornus and Alangium. Thus both wood anatomy and rbcL sequences support close affinity of Alangium, Cornus, Curtisia, Diplopanax, Mastixia, and Nyssaceae. Palynology also supports the above delimita­ tion for this family, there being a close relationship among Cornus, Afrocrania, Curtisia, and Mastixia, while Aucuba, Griselinia, Melanophylla, and Kaliphora are more dis­ tinct (Ferguson 1977). The placement of the other genera formerly included in Comales remains unre­ solved by wood anatomy. The analyses of rbcL sequence allied all these genera with a wide array of taxa in Asteridae s.l. (Xiang et al. 1993; Xiang & Soltis in press). Among them, Aucuba and Garrya form a clade si ster to Eucommia, and Griselinia, Aralidium, Toricellia, and Melanophylla are allied with representatives of Apiales. Corokia is in a clade with Argophyllum and two higher asterids, Helianthus and Lo­ belia, and Helwingia is allied with Phyllonoma and Ilex. The branchings among these genera in the present wood anatomical cladogram disagree with the above recogni­ tion of clades from rbcL sequences, and the further relationships of the taxa in clade 11 as suggested by wood anatomy clearly have to await a more general study. Hydrangeaceae and Loasaceae included in the Comaceous clade by the rbcL se­ quence analyses (Xiang et al. 1993; Xiang & Soltis in press) are not directly compa­ rable with the Comales. The latter are mostly arboreal while the secondary xylem of Saxifragaceae and Loasaceae shows much more derived characteristics, related to their shrubby or herbaceous habit. A cladistic analysis of wood anatomical data in­ cluding Deutzia, Philadelphus, Platycrater, and Hydrangea resulted in these taxa scattered in distant branches of the present cladogram: Hydrangea paniculata sister to Mastixia; Deutzia to Corokia; Philadelphus to Garrya; and Platycrater and Hy­ drangea-shrubs occupying several positions. This break-up of Hydrangea is due to its wide wood anatomical range which cannot be assessed properly with the character

Downloaded from Brill.com10/10/2021 08:45:33PM via free access Noshiro & Baas - Systematic wood anatomy of Comaceae 89 state allocation for Cornales. For example, some wood anatomical characters of Hy­ drangea (which includes shrubs, small trees, and lianas) vary almost as much as the whole order of the Cornales: vessel diameter ranges from 34 to 104 Iffil, bar number of perforation plate 10 to 50, rays all homocellular to all heterocellular, and ray height and width range from 0.094 to 18 mm and 19 to 164 f.lm, respectively (Stern 1978). The mainly herbaceous family Loasaceae is also specialised in wood anatomy and has 1) short vessel elements with simple perforations, 2) large rays consisting mostly of upright cells and some square cells (procumbent ray cells occur only in Mentzelia humilis), 3) scarcity or absence of uniseriate rays in several taxa (Carlquist 1984). Thus a totally different assessment of wood anatomical character states is necessary to clarify the phylogenetic relationship between these two families and Cornaceae. Inclusion of members of asterid clades of Chase et al. (1993), such as genera of Oleaceae, Araliaceae, or Pittosporaceae, may have clarified positions of genera in clade 11. However, the character scoring used for the present analysis of Cornaceae allies yielded too many multistate characters for some selected genera of the above families, and cladistic analysis in a broader circumscription was not feasible.

Wood anatomy and infrageneric classification The wood anatomical subdivisions within Cornus support monophyly of subg. Afrocrania, and subg. Mesomora together with Nyssaceae, and the positioning of subg. Syncarpea sister to this monophyletic clade (Fig. 82). This disagrees with both the results of the rbcL sequence analyses (Xiang et al. 1993; Xiang & Soltis in press) and the morphological phylogenetic analysis (MurrellI993). The rbcL sequence analy­ ses recognised three distinct clades: one consisting of subg. Syncarpea, Cynoxylon, and Arctocrania, one of subg. Cornus, and one of subg. Kraniopsis, Mesomora, and Yinquania. However, the relationships between these three clades are unresolved. The morphological analysis placed subg. Yinquania as sister to the whole genus and re­ garded subg. Mesomora and Kraniopsis to form a monophyletic clade sister to all the other subgenera, followed by the branching of subg. Arctocrania. The rest of the subgenera were divided into two clades, one of subg. Afrocrania and Cornus, and another of subg. Discocrania, Cynoxylon, and Syncarpea. The results of these two types of analysis closely agrees in the recognition of the basic three groups. If this is true, vessel grouping and bar number of scalariform perforation used for the subdivi­ sion within this genus should be taken to reflect parallelisms or reversals. The dia­ gram of Eyde (1988) was similar to the results of the rbcL data in the recognition of terminal clades and agrees with the morphological analysis in the basal placement of a clade consisting of subg. Kraniopsis, Mesomora, and Yinquania. Though subg. Yinquania is considered 'primitive' in the morphological analysis of Murrell (1993) and the systematic discussion of Eyde (1988), its wood structure is not much different from that of subg. Cornus, Cynoxylon, Discocrania, and Kraniopsis. In Alangium, sect. Rhytidandra forms a basal entity and differs from the other sections in vessel element lengths of 800-1600 Iffil and the absence of prismatic crys­ tals in axial parenchyma. The branching of the present cladogram differs from the schemes of Eyde (1968, 1972) using morphological and anatomical characters and

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Reitsma (1970) using pollen morphology. Eyde placed sect. Alangium and Constigma in one clade, and sect. Rhytidandra and Marlea in another, and Reitsma recognised three major branches from an ancestral fossil pollen type that did not correspond with the sections. This discrepancy partly reflects the ambiguous systematic position of Alangium grisolleoides, which has been allied with sections Constigma, Rhytidandra and Alangium (Capuron 1962; Eyde 1972, 1988; Govindarajalu 1979). Wood anatom­ ically, this species conforms to sect. Marlea in having scanty paratracheal parenchy­ ma, but the absence of prismatic crystals in rays and the length of vessel elements and fibres also agree with sect. Rhytidandra. Its vessel element and fibre lengths are out of place in both sect. Alangium and Constigma, and its simple perforation plates would be anomalous for sect. Constigma. However, only one specimen was available of A. grisolleoides. To clarify phylogeny within Alangium, further studies are necessary. In Nyssaceae, Nyssa javanica was distinct from the other Nyssa species in wood anatomy. The cladistic analysis of the morphological characters of Nyssa by Wen & Stuessy (1993) recognised three monophyletic clades, one of N. aquatica, one of N. javanica, N. shagszeensis, and N. talamancana, and one of N. ogeche, N. shweliensis, N. sinensis, and N. sylvatica. The present cladogram from wood anatomy agrees with this result in the recognition of a distinct lineage of N. javanica, but did not indicate a special position of N. aquatica.

CONCLUDING REMARKS

Wood anatomical diversity appears to lend itself weIl to contribute to OUf understand­ ing of the phylogenetic relationships within the order Cornales (sensu Cronquist). The high resolution of the cladistic analysis is surprising in view of the relative wood anatomical homogeneity and the limited number of cladistically useful characters. However, it should be stressed that several clades are only very weakly supported, and that only certain lineages are robust enough to attach great systematic value to them. The wood anatomical datamatrix presented here should ideally be incorporated in a future cladistic analysis combining morphological, palynological and molecular data (cf. Herendeen 1996 and Herendeen et al. in press).

ACKNOWLEDGEMENTS

We are espeeially grateful to Ms. I. Sakamaki and F. Endo for sectioning most sampIes; Ms. B.J. van Heuven, Dr. T. Fujii & Ms. Y. Kondo for technical assistance; to Dr. R. B. Miller and Ms. D. J. Christen sen, U.S. Forest Products Laboratory, Madison, U.S.A. for ample multiple supplies of wood speeimens; to Dr. W.c. Dickison, University of North Carolina, U.S.A. for a loan of Dr. J.E. Adams's materials; to Dr. S.R. Manchester, Florida Museum of Natural History, U.S.A., for the latest information on the systematics of North American Cornaceae and Nyssaceae; to Drs. Q.-Y. Xiang, Ohio State University, U.S.A., and D.E. Soltis, Washington State University, U.S.A. for allowing us to eite their unpublished paper; and to Dr. P. C. van WeIzen, Rijksherbarium, Leiden, The N etherlands, Prof. P. F. Stevens, Harvard University Herbaria, U. S. A., and Dr. P. S. Herendeen, The George Washington University, Washington, U.S.A. far critical reading and comments on cladis­ tics and phylogeny. For the generous supply of materials, we would like to thank Dr. H. Beeckman, Koninklijk Museum voar Midden Afrika, Belgium; Mr. P. D6tienne, CIRAD-Faret, Montpellier,

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France; Dr. P.E. Gasson, Royal Botanic Gardens, Kew, U.K.; Mr. LD. Gourlay, University of Oxford, U.K.; Dr. J. Ilic, Division of Forest Products, CSIRO, South Clayton, Australia; Dr. Yang Jiaju, Research Institute of Wood Industry, China; Dr. R.W. den Outer, Wageningen Agricultural University, The Netherlands; Dr. H.G. Richter, Federal Research Centre for Forestry and Forest Products, Hamburg, Germany; Dr. D. Tinambunan, Forest Products and Forestry Socio-Economics Research and Development Centre, Indonesia; Mr. W. Tze, Forest Research Centre, Sandakan, Malaysia; Mr. B.J.H. ter Welle, Utrecht University, The Netherlands; Ms. E. Wood and Dr. P.A. Groff, Harvard University Herbaria, U. S.A. Collection of several sampies was supported by the Monbusho International Scientific Research Pro gram (Field Research) (Nos. 58041022, 60041018, 63041060) from the Ministry of Education, Science and Culture, Japan.

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ApPENDIX - List of specimens studied

Abbreviations of wood collections follow the Index Xylariorum (w. L. Stern. 1988. IAWA Bull. n. s. 9: 203-252).

ALANGIACEAE Alangium sect. Alangium - A. longijlorum Merr.: Philippines, FHOw 12631, FPAw 23717, TWTw 2141. -A. salvijolium (L. f.) Wangerin: Hainan, China, CAFw 14610; India, FHOw 387 (A. larmirkii); Thailand, FPAw 27279, TWTw 1904. Alangium sect. Constigma -A. havilandii Bloemb.: Sarawak, FHOw 19297, TWTw 10525.­ A. javanicum (Blume) Wang.: Lematang Ilir, Sumatra, BZFw 30875; Java, FHOw 11942 (A. begoniaejolium); Malaysia, FHOw 12680, FHOw 12681; Kinabatangan, Sabah, SANw 98859; Sabah, TWTw 9513, TWTw 9514; Sarawak, TWTw 10526; Sula Archipelago, BZFw 29090; Philippines, TWTw 6575 (A. meyeri); Irian Jaya, BZFw 31189; Solomon Is., FHOw 18042. -A.javanicum var.papuanum Bloemb.: W New Guinea, BW 2922, BW 2958. -A. nobile (Clarke) Harms: Malaysia, FHOw 12683. - A. ridleyi King: Asahan, Sumatra, BZFw 6027; Kutai, Kalimantan, BZFw 33074, BZFw 33141; Malaysia, FHOw 12684. Alangium sect. Marlea -A. alpinum (C. B. Clarke) w.w. Smith & Cave: Dhawalagiri Zone, Ne­ pal, Ohba et al. 8340186; Bagmati Zone, Nepal, Suzuki & Noshiro 8540125; Sakuwasabha Distr., Nepal, Suzuki et al. 8840121. - A. chinense (Lour.) Harms: Burma, FHOw 3013 (Marlea begoniaejolia); Burma, FHOw 12621 (ditto); Burma, FHOw 12624 (ditto); Dhawaragiri Zone, Nepal, Suzuki et al. 8840509; Sikkim, India, TWTw 15376; Java, SJRw 22872; E Java, SJRw 31018 (M. begonifolia); Tokyo, Japan, TWTw 6374; Hainan, China, MADw 28039; Fujian, China, CAFw 9135; Guangdong, China, Tw 42061; Jiangxi, China, TWTw 7420; Tanzania, FHOw 5067 (A. begoniaejolium); Tanzania, FHOw 12629; Burundi, Tw 17222; Zaire, Tw 903; Forest ofNyungwe, Rwanda, Tw 28708. -A. grijfihii (Clarke) Harms: Singapore, FHOw 5764 (A. uniloculare); W Java, SJRw 31015 (M. densijlora); Sandakan, Sabah, SANw 81207. - A. grisolleoides R. Cap.: Perinet, Madagascar, CTFw 13802. -A. kurzii Craib: Sumatra, FHOw 12622; Yunnan, China, CAFw 3464 (A. tomentosa); Guangxi, China, CAFw 9268. - A. pla­ tanijolium (Sieb. & Zucc.) Harms var. trilobum (Miq.) Ohwi: Kanazawa, Japan, Kanazawa 119; Wakayama, Japan, TWTw 14174; Kyoto, Japan, TWTw 14816. -A.premnijolium Ohwi: Kyushu, Japan, TWTw 2228; Okinawa, Japan, TWTw 15199. - A. rotundijolium (Hassk.) Bloemb.: W Java, SJRw 31016 (M. tomentosa); Kota Kinabalu, Sabah, SANw 32381. Alangium sect. Rhytidandra - A. villosum (Blume) Wang.: W Java, SJRw 31011 (M. vitiensis); E Java, SJRw 31012 (ditto); Queensland, Australia, FHOw 12620, FHOw 12627 (A. vitiense); N. S.w., Australia, FPAw 10216; Wainimala Valley, Fiji, SJRw 37481 (A. villosum var. vitiense?).

CORNACEAE Aralidium - A. pinnatifidum (Jungh. & de Vriese) Miq.: Sumatra, FPAw 29006, MADw 27358; Java, SJRw 16054. Aucuba -A. chinensis Benth.: Sichuan, China, CAFw 151; China, SJRw 21794. -A. himalaica Hook. & Thoms.: Darjeeling, India, Kw 10749. -A.japonica Thunb.: Shizuoka, Japan, TWTw 5688, TWTw 6861; Tokyo, Japan (cult.), TWTw 6314. Cornus subg. Ajrocrania - C. volkensii Harms: East Africa, SJRw 3608, 27557. Cornus subg. Arctocrania - C. canadensis L.: Manza, Gunma, Japan, H. Hatta s. n. 7 Sep. 1977 & 6 Aug. 1979; Konsei Pass, Tochigi, Japan, H. Hatta s. n., 21 Aug. 1982. Cornus subg. Cornus - C. chinensis Wang.: Sichuan, China, CAFw 6169; China, SJRw 21808. - C. mas L.: N Greece, MADw 17876; Italy, MADw 26245, MADw 27924. - C. officinalis Sieb. & Zucc.: Korea, SJRw 32297; Ibaraki, Japan (cult.), TWTw 4299. - C. sessilis Torr.: California, USA, Kw 10787, MADw 27928. Cornus subg. Cynoxylon - C.florida L.: Tennessee, USA, MADw 8840; Louisiana, USA, MADw 8930; West Virginia, USA, MADw 23788. - C. nuttallii Audubon: Oregon, USA, MADw 2809, MADw 27926; California, USA, MADw 44696.

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Cornus subg. Discocrania - C. disciflora Moc. & Sesse ex DC.: Rio Macho Area, Costa Rica, MADw 24710; Chiriqui, Panama, MADw 24736; San Gerardo de Dota, Costa Riea, MADw 42286. Cornus subg. Kraniopsis - C. alba L.: Qingling, China, CAFw 5364; Brabant, Belgium, Tw 43798. - C. alsophila WW Smith: eult. at Kew (Siehuan, China) (branch), Kw 1988-8679. - C. amomum Mill.: Ohio, USA, MADw 9614; Wisconsin, USA, MADw 19206, MADw 22233.­ C. angustata: Hunan, China, CAFw 13429 (Dendrobenthamidia angustata). - C. asperijolia Michx.: Ohio, USA, SJRw 40652. - C. bretschneideri L. Henry: Anhui, China, CAFw 20632. - C. coreana Wang.: locality unknown, TWTw 12508. - C. darvasica Pojark: cult. at Kew (Tadjik, USSR) (branch), Kw 1993-1251. - C. drummondi Meyer: Missouri, USA, MADw 18300; Indiana, USA, MADw 18301. - C. excelsa H.B.K.: Chiapas, Mexico, MADw 23885. - C. glabrata Benth.: cuh. at Kew (W USA) (branch), Kw 1992-791. - C. macrophylla Wall.: Gandaki Zone, Nepal, Ohba et al. 8340106, 8340304; Suzuki et al. 8840442; Qingdao, China, CAFw 5745 (c. brachypoda); Siehuan, China, CAFw 19158; Shikoku, Japan, TWTw 910; Chiba, Japan, TWTw 13357 (c. brachypoda); Kagoshima, Japan, TWTw 13509. - C. meyeri (Pojark.) Pilipenko: cult. at Kew (Uzbek, Taskent) (braneh), Kw 1993-1250. - C. mombergii Hemsl.: Yunnan, China, CAFw 17256. - C. occidentalis (T. & G.) Coville: Wageningen, The Nether­ lands (eult.), PL 1535; Oregon, USA, Tw 45635. - C. parviflora Chien: Guangxi, China, CAFw 6421; China, SJRw 21905. - C. peruviana Maebr.: E Cordillera, Ecuador, FHOw 10987; Pichincha, Ecuador, MADw 16626. - C. poliophylla Schneid. & Wang.: eult. at Kew (braneh), Kw 1970-6180; Wageningen, The Netherlands (eult.), PL 1537. - C. purpusii Koehne: cuh. at Kew (Ontario, USA) (braneh), Kw 1987-645; Wiseonsin, USA, MADw 18076 (c. obliqua); Illinois, USA, MADw 18296 (c. obliqua). - C. racemosa Lam.: Ohio, USA, MADw 12193 (C.foemina); Indiana, USA, MADw 18297; Wisconsin, USA,MADw 22203.-C.rugosa Lam.: Wiseonsin, USA, MADw 8581, MADw 22115, MADw 25471. - C. sanguinea L.: ob Morell VS, Switzerland, Sehweingruber s.n.; England, FHOw 2207; Bagley, England, FHOw 2821.­ C. sanguinea L. subsp. australis (C.A. Meyer) Jar: euh. at Kew (W Asia) (branch), Kw 1993- 1256. - C. schinderi Wang.: eult. at Kew (Siehuan, China) (braneh), Kw 1994-1961. - C. stolonijera Michx.: California, USA, MADw 44613; Oregon, USA, MADw 44788; Washing­ ton, USA, MADw 45100. - C. stricta Lam.: Florida, USA, MADw 2408; Texas, USA, Tw 52620; Georgia, USA, MADw 21092 (C. foemina). - C. walteri Wang.: Sichuan, China, CAFw 8675; Korea, TWTw 7939.- C. wilsoniana Wang.: Guangdong, China, CAFw 17142, Tw 42059. Cornus subg. Mesomora - C. alternijolia L. f.: Georgia, USA, MADw 9741; Wiseonsin, USA, MADw 22113, MADw 25477. - C. controversa Hemsl. ex Prain: Hubei, China, CAFw 9844; Chiba, Japan, TWTw 13356; Kagoshima,Japan, TWTw 13510; Gifu, Japan, TWTw 13917 (Swida controversa) . Cornus subg. Syncarpea - C. capitata Wall.: Yunnan, China, CAFw 11906 (Dendrobenthamia capitata); Dhawalagiri Zone, Nepal, Ohba et al. 8340198, 8840536 (Benthamidia capitata); Sagarmata Zone, Nepal, Ohba et al. 8540266 (ditto). - C. ferruginea Wu: Guangdong, China, CAFw 16662 (Dendrobenthamiaferruginea). - C. hongkongensis Hemsl.: Guangxi, China, CAFw 15788 (Dendrobenthamia hongkongensis); China, SJRw 21937, SJRw 21971. - C. kousa Bürg. ex Miq.: Hubei, China, CAFw 9781 (Dendrobenthamia japonica var. chinensis); Kago­ shima, Japan, TWTw 13511; Gifu, Japan, TWTw 13932, TWTw 14829 (Benthamidiajaponica). - C. tonkinensis (Fang) Tardieu: Guizhou, China, CAFw 20289 (Dendrobenthamia tonkinensis). Cornus subg. Yinquania - C. oblonga Wall.: Tibet, China, CAFw 19267; Uttar Pradesh, India, Tw 45520. Corokia - C. buddleoides A. Cunn.: New Zealand, SJRw 25438. - C. collenettei Riley: Rapa, SJRw 37337, SJRw 37421 (Lautea collenettei). - C. cotoneaster Raul: cult. at Kew, Kw 1969- 18607. - C. macrocarpa Kirk: eult. at Kew, Kw 1986-797. - C. whiteana L. S. Smith: Queens­ land, Australia, FPAw 33986; New South Wales, Australia, SJRw 55149. Curtisia - C. dentata (Burm. f.) C.A. Sm.: Knysna, South Africa, MADw 16801 (C. faginea); SouthAfriea, MADw 28041 (ditto); Zimbabwe, MADw 28042 (ditto); Transvaa1, SouthAfrica, MADw 39527.

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Diplopanax -D. stachyanthus Hand.-Mazz.: Guangxi, China, CAFw 15828; China, USw 0042379 (branchwood). Griselinia - G. littoralis Raoul: Victoria, Australia, FPAw 10013. - G. lucida Forst. f.: Auckland, New Zealand, SJRw 25420; Patea, New Zealand, SJRw 50340. - G. ruscifolia Ball.: Cordil­ lera Pelada, Valdivia, Chile, BFHw 16776; Santa Catarina, Brazil, Uw 6953; Serrados Orgaos, Brazil, Uw 20826 (var. itatiaiae). Helwingia - H. himalaica Hook. f. & Thomas.: Koshi Zone, Nepal, Suzuki et al. 8840197. - H. japonica (Thunb.) Dietrich: Saitama, Japan, TWTw 944; Shizuoka, Japan, TWTw 14144; Kagoshima,Japan, TWTw 14374 (var. parviflora). Kaliphora - K. madagascariensis Hook. f. (branchwood): Angavokely, Madagascar, Bernardi 11058 (L); R. Brown s.n. (Paris 899). Mastixia subg. Manglesia - M. octandra Matthew: Mt. Kerintji, Sumatra, Meijer 6316. Mastixia subg. Mastixia sero Alternae - M. arborea (Wight) Bedd.: Mysore, India, MADw 33047; India, TWTw 11176. - M. cuspidata Blume: Lahad Datu, Sabah, SANw 132556; Gn Palung Nature Reserve, W Kalimantan, van Balgooy & van Setten 5471. - M. macrophylla (Thw.) Kosterm.: MasheliaDistr., Ceylon, Kostermans 27053, 27065. -M.pentandra Blume: Laguna, Philippines, MADw 28043. - M. pentandra subsp. cambodiana (Pierre) Matthew: Hainan, China, FRTGw 18 (M. alternifolia ). - M. pentandra subsp. philippinensis (Wang.) Matthew: Philippines, TWTw 5011, TWTw 11057 (M. philippinensis). - M. tetrandra (Thw.) Clarke: Ceylon, Kw 10796. -M. tetrandra var. thwaitesii Clarke: Hakgale Jungle, Ceylon, Kostermans 24187. -M. tetrapetala Merr.: Mt. Tabayoc, Luzon, Philippines, Jacobs 7518. Mastixia subg. Mastixia sero Oppositae -M. eugenioides Matthew: 30 km S ofMiri, NW Borneo, Fuchs 21311. -M. kaniensis (Melch.) Matthew subsp.ledermannii (Melch.) Matthew: C. 45 km N of Mindiptana, W New Guinea, TWTw 13806. - M. rostrata Blume: Atjeh, N Sumatra, de Wilde & de Wilde-Duyfjes 12714; W Java, SJRw 31022, SJRw 31023. - M. rostrata var. caudatifolia (Merr.) Matthew: Brit. N Bomeo, SJRw 20240 (M. caudatifolia). - M. trichotoma Blume: Atjeh, N Sumatra, TWTw 12674 (var. trichotoma); Atjeh, N Sumatra, de Wilde & de Wilde-Duyfjes 16702; Lahad Datu, Sabah, SANw 59252. - M. trichotoma var. maingayi (Clarke) Danser: Barisan Range, S Sumatra, de Vogel 1249. Melanophylla - M. capuronii Keraudren: Distr. Tamatave, Madagascar, CTFw 9052. - M. crenata Bak.: Distr. Fianasantsoa, Madagascar, CTFw 8879. Toricellia - T. arguta Oliv.: China, SJRw 21773. - T. tilii/olia DC.: Gandaki Zone, Nepal, Ohba et al. 8340334; Bagmati Zone, Nepal, Suzuki & Noshiro 8540005.

GARRYACEAE Garrya subg. Fadyenia - G./adyenii Hook.: Belmonte, Santa Clara, Cuba, SJRw 16712; Haiti, SJRw 19632. - G. glaberrima Wang.: Nuevo Leon, Mexico, Aw 33459. - G. lauri/olia Benth.: Michoacan near Morelia, Mexico, MADw 3777; Chiapas, Mexico, MADw 23809, MADw 23893. - G. longi/olia Rose: Mexico, MADw 22028, SJRw 53325. - G. ovata Benth.: New Mexico, USA, Tw 42478. - G. wrightii Torr.: New Mexico, USA, MADw 42082; Arizona, USA, SJRw 49475, SJRw 49478. Garrya subg. Garrya - G. buxifolia A. Gray: Oregon, USA, MADw 44892; Califomia, USA, SJRw 47259, SJRw 49471. - G. elliptica Douglas ex Lind!.: Oregon, USA, MADw 45068, MADw 45076; Califomia, USA, SJRw 47260. - G. .flavescens Watson: Arizona, USA, MADw 11278; Califomia, USA, SJRw 49482, SJRw 49485. - G./remontii Torr.: Oregon, USA, MADw 44842, MADw 44845; Califomia, USA, SJRw 47263. - G. veatchii Kellog: Califomia, USA, SJRw 49472, SJRw 49473, SJRw 49474.

NYSSACEAE Camptotheca - C. acuminata Decne.: Lu Shan, China, SJRw 21463; China, SJRw 21715; W Szechuan, China, SJRw 42577. Davidia -D. involucrata Baill.: Tokyo, Japan (cult.), Noshiro s.n.; Sichuan, China, TWTw 3194; China, TWTw 13651.

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Nyssa - N. aquatica Linn.: Mississippi, USA, MADw 1094; Louisiana, USA, MADw 8482; Texas, USA, SJR w 47159. - N. bijlora Wa1t.: F1orida, USA, Aw 28096; North Caro1ina, USA, MADw 8431 (N. sylvatica var. biflora); Georgia, USA, MADw 34424. -N.javanica (Blume) Wang.: India, FPAw 26001 (N. sessiliflora); W Benga1, India, Tw 17713 (ditto); N Thailand, Doi Sutep, TWTw 12529; Indonesia, TWTw 4454; Sumatra, BZFw 9999; Bengkulu, Sumatra, BZFw 17084; W Java, BZFw 20755. -N. ogeche Bartr. ex Marsh.: Georgia, USA, MADw 908; W Florida, USA, SJRw 40124; Georgia, USA, SJRw 48946. -N. sinensis Oliv.: Hunan, China, CAFw 9394; Guangdong, China, Tw 42065; Jiangxi, China, TWTw 7468. - N. sylvatica Marsh.: Mississippi, USA, MADw 1081; Missouri, USA, MADw 3027; Illinois, USA, MADw 15811. - N. ursina SmalI: Florida, USA, Aw 28094.

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