Wood Anatomy of Staphyleaceae: Ecology, Statistical Correlations, and Systematics

Flora (1985) 177: 195—216

SHERwIN CARLQuIsT and DAVID A. H0EKMAN

Rancho Santa Ann Botanic Garden and Pomona. College, USA

Summary Wood of Staphyleaceo.e is characterized by vessels which are mostly solitary; vessel elements are long with scalariform perforation plates (mostly more than 20 bars per plate) and with scalari. form, opposite, or alternate lateral wall pitting. imperforate tracheary elements range from fiber t-racheids with fully bordered pits somewhat less dense than those of tracheids (Euscophis, Sta phylea, Turpinio.) to fiber-tracheids with reduced borders on pits (Huertea) to libriform fibers (Tapiscia). Axial parenchyma is mostly abaxial, with tendencies towards vasicentric scanty and ray-adjacent cells and only a few diffuse cells. Rays are both inultiseriate and uniseriate. Hetero geneous Type hA. The multiseriate portion of multiseriate rays is often not sheathed with upright cells and consists of procumbent cells which often have bordered pits on radial walls. Rhomboidal crystals, tyloses, and dark-staining amorphous deposits are found in some but not all species. Quantitative features show wood of Stephyleo. to be less markedly mesomorphic than that of the other genera, a fact perhaps related to winter cold. The Mesomorphy index is held to be more useful in analysis of dicotyledon woods and in predicting relationship with ecology than a conduc tivity formula, because it runs parallel to ecological gradients, takes into account vessel element length (apparently related to embolism localization), and represents degrees of relinquishment of safety as woods become more mesomorphic. Statistical correlation among wood features of Sta phyleaceue show vessel element length related to imperforate tracheary element length and to ray height because all of those are linked to fusiform cambial initial length. Correlation between vessel -diameter and vessel element length is slightly weaker, as is inverse correlation between vessel diameter and vessel density (where packing constraints tend to enforce a correlation). The genera of Staphyleaceae can be easily characterized by wood anatomy features such as growth ring pre sence or absence, perforation plate bar number, presence of helical sculpture within vessels, type of imperforate tracheary element, presence of septa in fibers (in which case axial parenehyma is absent), ray dimensions, tylosis occurrence, and crystal presence. Wood of Staphyleaceae most closely resembles that of some Cunoniales (Saxifragales), and resembles that of Sapindales some what less. Introduction Wood anatomy of Staphyleaceae has been little studied, although descriptions of a few species may be found in various works (GREGuSS 1959; JANSS0NIus 1911; KRAUSE 1960; REcoRD & HEss 1943; S0LEREDER 1892); the summaries of SoLE REDER (1908) and METcALFE & CHALK (1950) are useful. Wood of Staphyleaceae is of potential interest because it is relatively primitive compared with that of niost dicotyledons. However, wood of Staphyleaceae proves to be more diverse than hitherto noted. Primitive woods tend to be confined to highly mesic situations. Does this correla tion (CARLQuIST 1975) hold true in Staphyleaceae? If so, what diversity in ecology occurs within the family and how is it related to wood anatomy? Other questions relative to wood anatomy of Staphyleaceae include those of phyto geography and phylogeny. Staphylea occurs in the Northern Hemisphere in eastern and western North America as well as in Europe and Asia. Euscaphis ranges from Korea and Japan to Taiwan and China. Huertea is found in Peru and the West Indies.

Stophleu

11.

Huertea

Euscuphjs Table

Species pnnOta

GRAy

trifolia CUATREcASAS

x R. holocarpa

colcinca Ga’s

bun’ilda

PAX

staph’iieon1e

gland

granadinn

elegans

& 1.

nis.

eubensis

niosa

Vvood

bolanderi

Srv. jupoeica

ZABEL

DC.

HEMsL.

characteristics

Carlquist

MADw-23753 Cariquist

MADw-8707 MADw-8195

Carlquist

MADw-3067

Carlquist

SJRw-37221

FPAw-DFP-29096 Carlquist

Carlquist

RSAw-ETHIMH

Car1quit

S.JRw-44481

F1’Aw-DFP-32998

SJRw-43415

MADw-5376

Evertt

PRFw-14690

SJRw-21955 MADw-5055

FPAw-DFP-13256

Collection

Cariqitist

1)w-35776

7269

15665

15661

15700

15660

15654

15663

15655

15683

Staphyleaceae

(RSA)

140 141

103

42 47

53 82 84

189

207 113

184

144

102

139

161

169

269

123

137

124

25 27

51 49 42

1,240

1,080

1,150

1,210

1,130

1,100

748

658

909

812

772

597

634 490

682 803

599

810 802

772 589

476

997

818

972

1.00

1.04

1.28

1.09

1.12

1.14

1.11 1.20

1.07

1.15

1.16

1.04

1.16

1.14

1.51

1.22

1.40

1.28

1.08

1.34 1.04

1.00

1.24 4

1.00

2.0

1.5 2.0

1.5

2.0

1.5 1.5 2.0

2.0 1.5 2.0

2.0

2.5 2.0 2.5

2.0 2.0

2.0

1.5

2.0

24 21

22 17

13 13

25 15

14 34 31

19 21

23 18

23 20

19 33

23 26

1,280

1,480

1,130

1,280 1,400

1,340 1,110

1,280

2,070

1,150

1,690 2,150

1,480 2,280

2,180

1,270

1260

1,980

1,930

937

930

993

891

896

843

5.0

5.5

4.5

4.0

5.5 4.0

5.5 4.0 4.0

4.0 4.5

3.5 4.5

4.5

4.5 4.5

5.0

4.0 3.5

2.5 5.0

3.0

3.5

4.5

5.0

1,340 1,080

1,070

1120

549

566

464

787

770

942

830 535 787

545

647

824 540

605 545

867 526

493

631

903 990

49 27

36 39

44 44

44 54

40 47

39 36

298

322

317

364

593

397 401

403

262

300 287

723

405 316 228

232

389 394 470

394

629 527

430

681

552

A. A,

A, A,

0 0

0 A,

1),

2,240

7,060 5,750

1,440

2,410

2,310

1,920

1,300

430 14

349

215

161

147

207

535

467

150

273

368

235

163

283

111 201

759

156.0

149.0

24.6

0.8

12.8

0.2 11.6

0.2

0.3

0.2

1.5

0.7

0.2

3.1

0.1

0.4

1.2

1.1

0.8

1.0

9.2

7.6 3.6 1.8

z 4 Tupiscuo sinensis SJRw-21767 59 86 698 1.40 2.0 7 25 1,240 2.5 321 36 276 0 478 1.4 liLly. Tuipiiiiu brechy- FPAw-DFP-30078 133 40 2,140 1.00 2.0 50 37 3,570 7.0 2,450 87 1,750 A, D, R 7,120 78.2 petuie LINDEN V

1’. ccernosa PRUUE SJRw-28643 101 10 887 1.48 3.0 27 32 1,650 4.5 1,180 51 888 A, D, H, 5,600 65.0 V T. heterophylle MADw-16548 84 41 1,100 1.12 2.0 29 29 2,200 3.0 1,050 64 578 A, V 2,270 12.0 (H. & P.) TOL, HSAw-MER-X8 113 20 1,480 1.40 2.0 31 40 2,270 3.5 1,140 47 610 A, V 8,450 82.3 XJRw.43279 88 54 1,200 1.56 2.0 31 26 2,070 4.0 1,030 29 574 A, V 1,940 11.0 2’. aepalensis WAI.I,. FPAw-DFP-12147 89 56 1,060 1.00 2.5 39 39 1,850 6.0 1,540 103 752 A, V 1,680 11.1 SJHw- 3881 81 58 1,180 1.08 3.)) 45 30 2,130 5.0 891 77 868 A, V 1,640 7.3 2’. nudifloru L. MADw-35779 96 33 1,310 1.00 3.0 30 20 2,350 6.0 1,730 82 709 A, V 3.770 25.3

2’. occidentalj MADw-23981 99 41 1,220 1.32 3.5 28 40 1,890 4.5 1,030 36 776 A, D, V 2,920 23.3 (Sw.) G. 1)ON MADw-24191 8<) 12 1,180 1.90 2.0 30 29 1,920 3.5 1,060 52 612 A, V 7,860 34.0 .JRw-54585 97 14 1,36)) 1.00 2.)) 41 14 1,980 3.)) 758 48 525 A, D, H, 9,420 63.2 V

2’. ova jolio Ean. FPAw-DFP-31887 99 5)1 1,430 1.20 2.0 42 38 2,740 4.0 1,070 56 912 A. B, V 2,520 17.2 MADw-18424 113 44 1,04)) 1.00 2.1) 45 41) 2,590 5.5 1,661) 113 998 A, V 4,180 36.8 RSAw-PFR1-518 112 35 1,430 1.04 2.5 42 38 2,790 5.0 1,355 8)) 700 A, V 4,550 44.7 2’. puniculatu VeNr. MADw-24387 139 14 1,540 1.4)1 1.5 27 38 1,932 2.5 2,14)) 102 1,170 A, V 15,900 277.0 ‘ 1. peiitaathu MAI)w-29l72 119 43 1,05)) 1.00 2.5 33 40 3,03<) 0.)) 2,040 1)12 1,08)) A, V 4,520 46.2 (un’.) LINDEN 2’. poniifera DC. SIAI)w-3578)) 155 33 1,40)) 1.0)) 2.5 27 37 2,530 0.)) 1,45)) 76 938 A, 1), V 0,521) 173.0 SFCw.1191-258 110 35 1,440 1.0)) 2.)) 3)) 34 2,480 5.0 1,510 71 1,03<) A, B, V 4,500 41.6 1’. sphaciocoipa $P’Cw-11575-137 177 18 1,7)1)) 1.16 3.)) 39 5)) 3,000 5.5 2,120 92 958 A, V 17,00)) 536.)) HAssa.

Legend for columns: 1, mean vessel (liaml

Tapiscia consists of a single species restricted to nlontane central and southern China. Turpinia is of unusual interest because it occurs in both the New World and the Old. In the Old World, TRrpinia ranges froni China and Japan to Taiwan, the Philippines, the indomalesiari archipelagoes. Thailand. Vietnam. Cambodia. southern India. and Sri Lanka. In the New World. Turpinia extends from southern Mexico through central America, the West indies. Colombia. Ecuador. to Peru (distributional data from KRAUsE 1960). In the habitats of the family as a whole. Staphyleaceae is located in notably mesic sites: shady valleys, cloud forests, along riverhottonis or streams. in the understory of tropical forest. Obviously these habitats are much more restricted than those of such families as Ericaceae or Rosaceae. The chief point of ecological diversity within the family very likely relates to whether freezing of an appreciable degree occurs in their habitats. Staphylea, Tapiscia, and, to a lesser extent, Euscaphis grow in climatic regimes with much more fluctuation of temperature than those which Huertea and Turp 1 nia experience. The relatively uniforni ecology of the family may make it a suitable one for studying what wood features are statistically interrelated with each other. Is there. for example. a correlation between vessel element length and ray dimensions? What. reasons can he advanced for various degrees of correlation which occur? The genera of Stctphyleaceae may be uniformly niesic, but they differ markedly from each other in such features as fruit type. Are similar discontinuities present in wood anatomy? The family has not always been constituted as it is at present. Tapiscia and Huertea have been placed in other families at various times (for a review, see KRAUsE 1960). Views on the affinity of the family seem to be in t.he process of change. The earlier view, that the family belongs in Sapindales, has been maintained by several authors (CR0NQUIsT 1981; IJAHLGREN 1980: TAKHTAJAN 1980). This relationship was claimed because of such features as the pentamerous flowers, alternate leaves, and follicular fruits (Euscaphis, Staphylea). The inflated fruits of iS1taphylea simulate those of Cardiospermum or Koelreuteria. However, these appealing similarities may he mis leading. Since early in the twentieth century. some authors have believed Staphytea eeae should he placed in Saxifragales (Cunoniales). This view has been reviewed by DECKISON (1980): a recent advocate is THORNE (1983). Wood anatonw is significant in this regard because Staphyleaceae have markedly more primitive wood than do most Sapindales. Materials and Methods

All wood specimens were avai]ab]e as dried samples. The writer collected wood samples of various species of Staphylea in European botanic gardens in 1981. The kindness of the directors of the botanical gardens of Copenhagen and Goteborg is acknowledged. Tn 1982, collections were made of wood of Euscaphis japonico and Staphylea bum cildu in Japan. This study could not have been pursued without a large number of wood samples from arious xvlaria. Foremost among these is the U.S. Forest Products Laboratory, where Dr. Rrars B. MILLER was generous in sharing wood samples from the MADw and SJRw collections there. ‘I’lie Forestry (‘ommission of New South Wales offered samples (SF(’ numbers in Table 1). J. D. BRA ZIER of the Princes Risborougli Laboratory furnished other sample. The species Eiiscaphis sic phyleoi’lee S. & Z. may he conspecifie with E. jopoaica Pix but is recognized here for mainland Asiatic populations. 1-Ierbarium vouchers for my- own collections are located in the herbarium of the Ranchio Santa Ann Botanic Garden. Woods of some Stophyleaceae were sectioned on a sliding microtome after they were boiled. For others, an alternative method (CARTQI tsr 1982 a) was applied. The latter method uas not entirely successful, for woods of Stophyleaceae are only moderately soft, and most are probably host sectioned on a sliding inicrotome despite scattered instances of wall fracturing. sect ions were stained with safranin ; eounterst.aining with fast green aided iii demarcation of pit cavities. Mace- rations were prepared W itli JEFFREY’S Fluid and stained with safranin. Wood Anatomy of Stophyleoceoe 199

Table 2. Pearson’s Correlation Coefficients for wood features of Staphyleaceae (lower triangle = I-i; upper triangle = 112).

VL VI) VM VG BAR FEEL ITED 5TH MW tTH

VL —— 0.70 0.55 s 0.40 0.91) 0.49 0.07 * 0.75

VD 0.84 —- 0.73 * * 0.00 0.55 0.43 * 0.52

VM -—-0.74 —0.86 -— * * 0.51 * * * 0.38 VG * * * — * * * * * 0

lIAR 0.68 * * * — 0.49 * 0.56 * 0.4$

ITEL 0.95 l).81 -——0.72 * 0.70 —— 0.47 0.64 0.42 0.69

PEEl) 0.70 0.74 * * * 0.69 --— 0.39 * 0.47

MH 0.82 0.66 * * 0.75 0.80 0.li2 —--- 0.53 0.81

MW * * * * * 0.64 * 0.73 -— 0.44 UH 0.87 0.72 —0.61 * 0.69 0.83 0.68 0.90 0.66

Legend for columns: VL = vessel element length; VI) vessel diameter; XTM = number of vessels per mm2; VO = nimiher of vessels per group; BAR = number of bars per perforation plate; I’I’EL = imperforate traeheary element length; ITEI) = iinperforate traeheary element diameter; MR

multiserate ray height; MW niultiseriate ray width; RH = uniseriate ray height. It values felow 0.60 are represented by *.

Data in Table 1 are based upon 25 measurements per feature, except for imperforate tracljeary element diameter and wall thickness and vessel element will thickness, where a typical conditiou has been recorded. Vessel diameter includes the wall, as is usually done, although without doubt when considering the functional nature of the vessel one should consider the lumen only, as is done by OEvEa at al. (1981). However, those authors show that statistically the two measurements are virtually interchangeable. Data on wood anatomy of Staphyleccene have been subjected to a statistical analysis, that of PEARsoN’s correlation coefficient, in Table 2. Values below 0.60 have been emitted from Table 2 because in biology they are less significant. R values in a tabular form would occupy unIv half of the table, or one of the two triangles of the table. The H values given represent natural log transformations of the data, a procedure recommended when a relatively small number of data points are available. In the other triangle of Table 2, H2 values are shown. H2 values repre sent the proportion of the variance which is explained. Sections of the majority of species were prepared by the junior anther. Data were collected by him and by Dr. Dxvrn C. MIcn:ENES. Photographs and text construction are the work of the senior author.

Anatomical Descriptions

Growth Rings Huertea and Turpinia lack growth rings. The sample of Tapiscia was not sufficient to demonstrate nature of growth rings, although the last-formed vessels just beneath the cambium were narrower and may have represented the beginning of latewood. in Euscaphis, earlywood of growth rings contains wide vessels and wide fiher-tracheids. somewhat thinner walled than those of latewood fiher-tracheids (Fig. 1). Ray cells are radially narrow at the junction between latewood and earlywood. The growth rings of Staphytea are similar (Figs. 12, 14, 17), and somewhat more sharply demarcated than those of Euscaphis. Tessel Elements Vessel diameter is shown in Table 1, column 1. if one examines the values, one finds marked differences among the genera. The mean vessel diameter of Euscaphis (Fig. 1) is 73.2 sum. A lower mean vessel diameter is registered for the genus Staphytea

—______All

Turpinia. in

predominate to tends vessel-ray pitting Scalariform heterophyila. T.

except

Turpin’ia of

species all

and .inensis, Tapiscia L.. trifolia •S’. L., pinnata S.

x elegans

ZAREL,

S. S’cEv.,

colchica Staphylea granadina. H. GRISEB.. cubensis Huertea

staphyleoides,

Euscaphis in

types, two those to addition in noted, pitting riforni

scala species; most in

alternate

opposite is pitting Vessel-ray pitting.

vessel-vessel

and.

pitting vessel-ray of

terms

in discussed he can vessels of wall pitting Lateral

32.

and 26. 16. 15. 6, 11. Figs. by comparing seen he can orders of

presence

Relative 32).

Fig. (e.g..

ends their at appearance marked a of borders hear

Bars bordered.

usually are

wide) 3 4 um (about bars thicker whereas extent, ciable

appre any t.o

bordered not are bars wide) l 4 um (about the slenderer general, In DC.

era T. pomif and

icuiata VENT.., pan T. 32). (Fig. ELM. ovalifolia T. G. Don. (Sw.) tails

occiden T.

WALL., nepalensis T.

P.) TuL., & (R. heterophylla T. SPRucE. carno.sa T.

2—3um wide

hut

wide. 1—2um mostly are bars Turpinia, Tn 26). (Fig. wide

2—3 um

are Tapiscia

in Bars latewood. in wide 2—3utn are they 15): (Fig. vessels

earlywood

wide 1—2um

from range bars In Staphylea. 11). (Fig. wide 3.5uni and

between mostly

wider, appreciably are they Huertea. wide; in 1—2uni 6), (Fig. thin

bars relatively

are In Euscaphis, present. is bars of thickness in Variation phyleaceae.

Sta

observed were

plates perforation No simple infrequent. are 16) (Fig. plates

aberrant or other foraminate

genera: these of in all he forked may bars Occasional

32). Turpinia (Fig.

35 in

6) and (Fig. Enscaphis 31 in 15). (Fig. Staphylea in 21.6 11),

Huertea (Fig. in

14.3 to 26)

(Fig. Tapiscia in plate bars per 7.4 from range genera the

for

6. Means

column 1.

Table in species l)late each for shown are perforation per bars

number

the

for

Means 16). (Fig. thereof modification sonic or :32) 26, 6. 11, (Figs.

scalariform

as exclusively described he can Staphyleaceae in plates Perforation

2). (Table relation

cor

significant statistically

a not is there thickness, wall vessel and diameter vessel

between

parallel

notes one Although 2.37 Ia. Turpin viii. 2.OQum: Ta.pisc’ia. unì: 1.84

Staphylea. 1 tm; 2.25

Huertea, i.80um; Euscaphis. are: feature this for genera the for

Means

family. the

within little varies also 4) column 1, (Table thickness wall Vessel

1984). (CARLQuIsT dicotyledons of groups other many with compared

grouping of

vessel

degree appreciable an represents these of None Turpinia. in 1.23 and

in Huertea,

1.40

Staphylea. in 1.12 Tapiscia. in 1.38 Euscaphis. in 1.07 are genus h

Means CuATRECAsAS.

granadina Huertea in 1.51 is high this of Turpinia; species six

japonica and

in Euscaphis 1.00 is reported low The family. the throughout uniform

relatively is

feature This etc.). 2,

vessels of a pair 1. vessel solitary a = calculation,

this

(in 4

column 1, Table in calculated been has group per of vessels number The

1.390um. is genus the for mean the in which 28), (Fig. Turpinia in longer

much are elements Vessel 1,110,um. average 1) (Fig. 1,0301m1; Euscaphis of those

averaging elements vessel 8) has (Fig.

Huertea 697 m. 13. 18). (Figs. S’taphylea and

Inn,

683 24),

(Fig.

Tajiscia in found is figure lowest the genus. by genus calculated are

means if

family. the in great relatively 3) is column 1. (Table length element Vessel

35.0. with

28), (Fig.

Turpinia and of 37.0 mean a with 7). (Fig. Huertea are dimension this for

Lowest 1),

52.3.

(Fig. Euscaphis or 59.1. 23). (Fig. Tapiscia for mean the double than

more is figure

That nim 2 . per vessels of 148 number mean has a whole a as genus the

which in

Staphylea, in reached is A high AS’taphyleaceae. in density vessel for values

range

wide a moderately finds one 2. 1, column Table examines one if 2). (Table

correlated

highly

not is this although diameter, vessel to relationship inverse an bear to

tends

density) vessel termed (also transection of 11n1 2 per vessels if number The

urn. 110

is genus

the for mean the and ,urn, 177 80 to from ranges diameter which in

28), (Fig. Turpinia in

are clearly vessels widest urn. The 86.3 diameter, intermediate

vessels

has

23) (Fig. Oliv. sinensis

Tapiscia 49.8um. 17). 12. (Figs. whole. a as

H0EKMAN A. D. and Sir. CAp,LQuIsT 200 JH1 Wood Anatomy of Stephyleaccac 201.

Figs. 1—6. Wood sections of Euscophis joponiea (CARLQUIsT 15683). 1. Transection; earlywooci above center. 2. Tangential section; wide multiseriate rays contrast with uniseriate rays. 3. Radial section; wing of multiseclate ray above, procumbent cells of central portion below. 4. Portion of fiber-tracheid from tangential section, showing pits. 5. Portion of vessel wall from radial section; helical sculpture lies between pits. 6. Portion of scalariform perforation plate from radial section. Figs. 1—2 magnification scale above Fig. 1 (finest divisions = lOfLm). Fig. 3, scale above Fig. 3 (divisions = 10 tm). Fig. 4—6, scale above Fig. 4 (divisions 10 tim). 202 Sn. CARLQuIsT and B. A. H0EKnAN

Figs. 7—11. Wood sections of Huertea. 7, 9—11. H. cubensis (MADw-5376). 7. Transection; vessels wide, occasionally in pairs. 8. H. grenadine (MADw.25776), tangential section; multi senate rays short, various in width. 9. Subdivided erect ray cells bearing crystals from wing of iav as seen in radial section. O. Chambered crystals in fiber from radial section. 11. i’ortion of perforation plate from radial section. Figs. 7—8, scale above Fig. 1. Figs. 9—-] 1, scale above Fig. 4. Wood Anatomy of Stephyleaceae 203

Figs. 12—16. Wood sections of ,Stuphyleu boloiidrri (MADw-3067). 2. Transection; margins of two growth rings evi(lent. 13. Tangential section; ray widths vary. 14. Radial section through ray (wing at top); narrow cells (left) demarcate growth ring. 15. Short perforation plate from radial section. 16. Foraminate perforation plate from radial section. Figs. 12—-13, scale above Fig. 1. Fig. 14, scale above Fig. 3. Figs. 15—16, eale above Fig. 4.

Fig. above

scale

19—22, 1.

Figs. Fig. above scale 17----.18, Figs. pattern. like

tracheid showing

section,

radial from

elemont

tracheary Imperforate 22. evident. sculpture helical

section; radial

from wall

Vessel 21.

15655).

(CARLQUI5T 5. holocarpa 21—22. section.

radial

from

ccli

ray

Perforated 20.

section.

radial

from tylosis

thick-walled Relatively 19. wings.

uniseriate

tall

bear rays

multiseriate most

section;

18.

Tangential illustrated. rings growth three of margins

Transection; 17.

15654).

S. (CARLQUIsT bumaicla

17—20. of Staphylea. sections Wood 17—22. Figs.

A. D. and HoE1crAN Ss. CARLQUIsT 204 Wood Anatomy of Staphyleaceae 205

Figs. 23---—27. Wood sections of Tupi8cio sinensis (SJRw-21767). 23. TTansection; libriform fibers thin-walled. 24. Tangential section; rays short, septa evident in libriform fibers. 25. Radial section, showing vessel-ray pitting above and procumbent cells below. 26. Scalariform perforation plate from radisi section; bars wide. 27. Opposite and alternate pitting on vessel wall from radial sec tion. Figs. 23 24, scale above Fig. 1. Fig. 25, scale avove Fig. 3. Figs. 26——-27, scale above Fig. 4.

14 I’lora, Bd. 177 206 SR. CARLQU1ST and D. A. Ho1rMANN

Figs. 28—32. Wood sections of Terpiiiiu. 28 30. ‘P. occideutolis (MAI)w-24191). 28. ‘lianseetion; vessels few, fiber-tracheids wide. 29. Tangential section; rays tall. 30. Subdivided erect ray cells from wing of ray in radial section; each cell bears a crystal. 31 32. 1’. ovulifolie (RSAw-PFRT-518). 31. Bordered pits on fiber.traheids from radial section. 32. Portion of perforation plate from radial section; borders near ends of perforations. Figs. 28-29, scale above Fig. 1. Fig. 3() 32, scale above Fig. 4. Wood Anatomy of 3taphyleaceae 207

Measured vertically, vessel-ray pits are 5—--6 ni in Euscaphis and Staphylea, about 9um in Huertea (Fig. 11), 5-—7um in most species of Turpinia but 8—9,um in T. carnosa SPRucE, T. heterophylla, and T. pornifera. Scalariforni vessel-vessel pitting can he reported for Huertea cubes ,sis, H granadiiue, S’taphylea colchica, S pinnata, Turhinia hrachypetaia (ScHLECHTFR) v. P. LINDEN. T. carno.a, T. heterophylla, T. ‘nepalensis, T. nudifiora L., T. ovaiifoiia, T. pan iculata. T. pentandra (ScRLEcHTER) V. D. LINDEN, and T.sphaerocarpa HASSK. However, opposite pits (Figs. 5, 27) can also he found in .Euscaphis, Huertea, Staphylea (except S. bolanderi and S. bumalda), Tapiscia sinensis, Turpinia brachypetala, T. heterophylla, T. nepalensis, T. occiden tales, T. pentandra, and T. pomifera. Alternate pitting (an be found in Euscaphis japon.ica, E. .staphyleoide, Staphylea bolanderi, S. bumalda, S. x elegans, S. pinnata, S. tr ijiia, Tapiscia sinen.ce.s. Turpinia heterophylla, T. nudiflora, and T. occidentaiis. The three pitting types intergrade. Vertical dimensions of vessel-vessel pits are in uch like those given above for vessel-ray pits. Some pits with irregular pit apertures appearing somewhat like pit vestures were observed in Huertea glandulosa R. & P. This appearance may be due to droplet-like deposits. Helical sculpturing occurs in vessels only in the genera Euscaphis and Staphylea, but it was observed in all species of these genera. When least conspicuous, as in Sta phylea holocarpa HEMSL. (Fig. 21), helical sculpture appears as a pair of wisplike bands near each pit. fading out away from the pit. Where more conspicuous. as in K. cot- chico. the thickening bands interconnect pits. Euscaphis (Fig. 5) shows both of these types. The tYpes intergrade. and in no case are clear continuous helices of wall material present. The drawings of GREGuSs (1959) for S. coichica and S pin.nata are therefore oversimplifications. In Euscaphis and Staphylea, the helical sculpture is least visible in earlywood vessels, most conspicuous in latewood vessels, and was not observed in carlywood of B. japonica and S. bolanderi. Helices were observed on most vessels (as well as in some fiber-tracheids) of Euscaphis staphyleoi€les.

Tm perforate Traehear elements Staphyleaceac are unusual for a small family in that iniperforate tracheary eleimicnts range from fiber-traeheids which are very nearly tracheids (and could arguably be termed. so) to fiher-tracheids with variously reduced pits and even libriforni fibers (Tapiscia only) with simple pits. In the libriform fibers of Tapiscia sinensis. the pits are 2—3 iim in length (Fig. 26, left). In Euscaphis. iiiiperforate. tracheary elements bear fully bordered pits 5—6piii in diameter (Fig. 4). Sparseness of pits suggests these cells should he termed fiber-tracheids. In Staph.yiea. pits are also fully bordered and 5—6pnm in diameter (Fig. 22). The density of pits on. at least sonic imperforate tracheary elements of Staphylea leads one to term such cells tracheids, although fiber tracheids in which pits are slightly less dense can also be found in Staphylea woods, intergrading imperceptibly. The figure of GREGuSS (1959) in which imnpcrforate t.raehear elements with simple pits are illustrated for S. coichica umay represent diffi culty in observing borders. 1 found all iniperforate t•raeheary elements in mv material of that species bore bordered pits. In Turpisia, pits on imperforate tracheary elements are all fully bordered (Fig. 31). Diameter of these bordered pits is about the same as those. on vessels in Turpinia species respectively (for size, see account on vessel-vessel pits above). In some species of Turpinia, pit apertures are longer than the circular pit cavity outline. These elon gate, slitlike apertures umay owe part of their length to splitting of a mild sort in the wall, a splitting resultant. from dehydration. in .Huertea, pit borders of iniperforate t.raeheary elements are reduced, so that. the circular outline of the pit cavity is only 2—3 nm in diameter. The slitlike pit aper 14* 208 Sa. CARLQUI5T and I). A. H0EICMAN

tures in these cells extend beyond the circular outline of the pit cavity. Thus Huertea has fiber-tracheids approximately intermediate in pit features between tracheids and libriforni fibers. Vasicentric tracheids have been reported for Staphyleaceae (e.g., METCALFE & CHALK 1950). GREauss (1959) reports both tracheids and fiber- tracheids for Staphylea coichica and AS’. pinnata. HEIMScH (1942) also uses these two terms for iniperforate trachearv eleitients of Staphyleaceae. However. HEII.scH uses the terms tracheid and fiher-tracheid in the sense of BAILEY (1936). as I do: a fiber-tracheid is considered an imperforate tracheary element in which pits have smaller borders that those on tracheids, or the pits themselves, if fully bordered, are smaller or more sparsely distributed than those typical of tracheids. My adoption of this definition is aided by the apparent inefficiency of fiber-tracheids at conduction, as indicated by vessel grouping patterns (CARLQuIST 1984) in dicotyledons. METCALFE & CHALK (1950) appear to base their report of vasicentric tracheids in Staphyleaceae on the account of HEIMscH. and may represent to some degree a misinterpretation of HEIMsCR’S terms (ME’rcALFE & CHALK usually consider “ordinary” tracheids to be “fibres with abundant bordered pits”). To be sure, occasional imperforate tracheary elements have helical thickenings, like those of vessels, in the genera Euscaphis (HETMScH 1942) and Staphylea (GREauss 1959). Disregarding the helical thickenings in imper forate tracheary elements, I was not able to delineate two classes of imperforate tracheary elements within any given wood sample of Euscaphis or Staphylea. Sonic of the imperforate tracheary elements in these species have slightly arger pits than do others, but differences appeared minor, with no distinctions possible among such cells. In my material of Euscaphis staphyleoides, nearly all imperforate trac-heary elements hear helical sculpture, whereas few do so in the other Euscaphis collections. Terming these sculptured fiher-tracheids as vasicentric tracheids in B. stciphyleoides despite their identity with fiher-tracheids in E. aponica except for sculpturing seems illogical. Length of imperforat.e tracheary elements in Staph yleaceae (Table 1, column 8) parallels that of the vessel elements. Means for the genera are: Euscaphis, 2,100,um; Huertea. 1,63Quni; Staphylea. 1.140,am; Tapiscia, i.240,um; Turpinia. 2,370/Lfll. If one divides imperforate tracheary elements by vessel efment length, one obtains; the following figures: Euscaphis, 1.89; Huertea, 1.58; Staphylea, 1.64; Tapiscia, 1.82; Turpinia, 1.70. Imperforate tracheary element diameter (Table 1, column 7) is greatest in Turpinia (genus mean, 34.3 gin), least in Staphylea genus niean, 20.6 ni). Imperforate tracheary element wall thickness (Table 1, column 9) is not directly related to cell dimensions. hut is greatest in Turpinia. However, sonic species of Turpinia have notably thin fiber-tracheid walls (Fig. 28). The thinnest imperforate trachearv elements walls observed in the family were those of Tapiscia sinensis (Fig. 23). Wall thickness of fiher-tracheids varies with relation to position within growth rings in Euscaphis (Fig. 1) and Staphylea (Figs. 12. 17). Sept-ate nature of imperforate tracheary elements was observed in all species of Hnertea and Tapiscia (Fig. 24). In both genera, 3—4 septa per cell are typical. Iiiiper forate tracheary elements in the other genera were not observed to be septate.

Axial Parenchyma Distributions of axial parenchyuia are showii in Table 1, column 13. In Huertea (Figs. 7, 8), axial parenchyma was observed to be absent: this is true in Tapiscia (Fig. 23) also. In the other genera, abaxial parenchyma can be said to be present, but one or more other types may be observed also. Ahaxial plus some vasicentric cells Wood Anatomy of Staphyteea 2.09

can be seen in Euscaphis japonica (Fig. 1) relatively clearly contrasted with fiber tracheids in that photograph because of the thick-walled nature of fiber-tracheids. The combination of abaxial plus vasicentric scanty parenchyma is a common one in the genera other than Huertea. Diffuse parenchyma cells are also present, but to a limited extent, in an appreciable iiiimber of species. In addition, a few species showed a parenchyma type which has not hitherto been noted in dicotyledons. I am terming this type “ray adjacent” because parenchyma cells occur along the sides of rays (e.g., Fig. 1, right side of large ray, center). These cells may he regarded as a modification of a diffuse pattern in which the proportion of axial parenchyma cells in contact with rays is much higher than what one would expect in a random situation. Axial parenchyma strands consist of cells the number of which seems related to length of fusiforni cambial initials. For example, in Euscaphis japonica, a strand cell number of 4, 5, or 6 is typical, hut in Turpinia ovalifolia, the number ranges from 6 to 15 and typically is about 8. A feature atypical for strands of axial parenchyma cells is the tendency in Staphyleaceae for occasional walls to he markedly oblique rather than transverse. This tendency was noted in Eu.scap hi8 japonica (Fig. 2, middle left) and Staphylea trifolia. Vascular Rays Rays in Staphyleaceae are both multiseriate and uniseriate, the difference some times marked (e.g., few biseriat.e or triseriate rays, only wider rays present) and thus suggesting what some authors term ‘rays of two distinct size classes.” Uniseriate rays are more frequent than multiseriate rays in any given sample (Figs. 2, 8, 13, 29) or, in Tapiscia, somewhat less frequent (Fig. 24). The ray type of the family corre sponds approximately to Heterogeneous Type hA, of KRIs (1935). The rays of iluertea come closest to the description of KRIBS because erect cells tend to sheathe parts of the niult senate portions of inultiseriate rays (Fig. 8). The rays of Euscaphis (Fig. 2), Staphylea (Figs. 13, 18), Tapiscia (Fig. 24), and Turpinia (Fig. 29) tend to lack erect cells on the sides of niultiseriate portions of multiseriate rays. The marked differences between procumbent cells in the center of rays and upright cells in the iiniseriate wings of multiseriate rays can he seen in the radial sections of Figs. 3, 14, and 25. To be sure. some of the cells which appear in tangential section as though they might he upright, cells are, in fact. slightly longer radially than vertically (Fig. 3, center). However, such cells are nearly square as seen in radial section and are mar kedlv different from the procunihert cells. A multiseriate ray may contain a vertical strip of radially short cells, which must he termed square or erect, where growth slows at the end of a growth ring (Fig. 14). Otherwise dimorphism between wing cells arid multiseriate portion cells in multiseriate rays is clear. The uniseriate rays of Staphyleaceae are composed wholly of erect cells. Height of rays is shown in Table 1 (multiseriates, column 10; uniseriates, column 12). These heights parallel lengths of vessel elements of imperforate tracheary elements. Multiseriate rays average about twice as tall as uniseriate rays. Tapiscia sinensis is an exception to this: rays in this species are notably short.. Width of multiseriate rays is skown in Table 1, column 11. Notably wide rays are seen in Euscaphis japonica (Fig. 2) and Staphylea bolaruferi (Fig. 13); narrower rays are illustrated for Huertea cithensis (Fig. 8), Staphylea bumalda (Fig. 18), Tapiscia •inensis (Fig. 24), and Tur pinia occidentalis (Fig. 29). Perforated ray cells were observed in Staphylea bunialda (Fig. 20). All species of AS’taphyleaceae may be said to have ray cell walls which are moderately thin, hut which are lignified. Simple pits occur on radially-oriented walls. However, if one studies radial sections, one finds that tangentiallv-rtinning ray cell walls often bear bordered pits.

habitat the describe not does (1981) OLIvER Although specimens. Staphylea some

range in the lying family, the for (478) value Mesomorphy a has low Tapiscia

deciduous). is (Staphylea Euscaphis of habit evergreen

and

values Mesomorphv higher the in reflected fact is This unlikely. quite freezing

plant) (or soil of chance the renders influence maritime some localities, these in Even

China. of parts montane and Japan. southern Korea. southernmost in those are grows

Euscaphis which in areas coolest The Staphylea. of those than higher appreciably

are (759—2410) Buscaphis of values Mesoniorphv The E’uscaphis. of those than cated

demar sharply more Staphylea are in rings growth the However, habitats. seasonal

occupY

to be can said both that .so rings growth ylea Staph have and Euscaphis Both

unavailable). is and water frozen is the ground where foliage by transpiration (e.g..

drought physiological produces Freezing winter. in occurs regularly freezing where

habitats, seasonal strongly in lives it that fact due the to he may values is Mesoniorph

Staphyleaceae Stapht,’lea other the below falls that fact The places. wet permanently

and similarly bottoms stream to inhabit Staphylea of tendency the betokeris this and

minimum, at even mesic is still (147—535) genus this in range The 600. below fall all

Staphylea collections the that finds one values, Mesomorphy the one If analyzes

(1981). al. of et that OEvER notably papers. few a in only

used been

has figure this 15). although column 1. (Table Conductance figure the for

shown is The same 1981). CARLQusT (e.g.. Pittosporaceae: represented is Staphyleaceae

occupied than of habitats range wider a which in families those in species xeric

with compared as range a niesic in fall species all hut 14). column 1, (Table values of

range a one. finds 1977) (CARLQUIST Mesomorphv termed ratio the one calculates if

Conclusions Ecological

of occurrence. mode of listings above two

the in included not taxa the in observed not were deposits Amorphous occzcentatjs.

Turpinia and trifolia. S. holocarpa, S. S. bumaida. bolanderi. Staphylea in served

ob were droplets of form the in deposits limited More porn ifera. T. and pentandra,

T. ovalifolia, T. T. nudiflora. nepalensis, T. heterophylia. T. carnosa. Turpinia 25).

24. (Figs. .sinensis Tapiscia Huertea. of species 9) other 8 and (Figs. cuhensis Huertea

(Fig. japonica Euscaphis of ray cells in he massive to observed were deposits These

Staphyleaceae. most in observed were nature) chemica. in not probably (but texture

in resinlike appear and which amorphous are which of materials Accumulations

Turpinia. in observed otherwise not

were 30): crystals (Fig. is occide??tal T. species of the collection one only in observed

were crystals hearing cells ray erect Subdivided 10). (Fig. crystal rhomboidal large

single a bearing each cells, numerous into septate are fiber-tracheids sonic cubensis

Huertea in In addition, cells. right) 9. upper three (Fig. rarely or two into divisions

sub these from resulting cells of the in each occurs crystal rhomboidal One large

9). (Fig. studied species of all cells ray erect in subdivided observed were crystals

Huertea, In iscia. Tap or ytea. Staph Euscaphis. in observed not were Crystals

inclusions Cell

T. pomifera. and pentandi’a. T. l. paniculata, lensis,

uepa T. brachypetala, T. species: some in only observed were tyloses In Turp’inia,

19). (Fig. burnaWa Staphylea for illustrated are walls thicker Slightly 24). (Fig. cia

Tapi in as vessels filling sometimes thin walled, very mostly were These found. were

tyloses

genera, four other the in Euscaph’i$: in the observed genus not were Tyloses

Tv loses

A. Hoz D. and IcRAN SR. CARLQUI5T 210 Wood Anatomy of Staphyieaceoe 211

Tapiscia sinensis, he does mention presence of winter buds, suggesting the plant is deciduous and that it may occupy a climate with cold winters like those which prevail where Staphylea grows. Ruertea and Turpinia have uniformly high Mesomorphy values. These species occupy frost free areas which are perpetually moist. Although Huertea and Turpinia tend to be understory elements, their relatively high transpirational characteristics (suggested by high Mesomorphy and Conductance values) may he related to their broadleaved evergreen habit and by their occupation of warm areas. Within genera of Staphyleaceae, one cannot discriminate among species effectively as to which has a more or less wet habitat: all st.aphyleaceous habitats may he de scribed as wet. Variation in Mesoniorphv can occur because of age of tree, since most trees tend to develop wider and longer vessel elements with age, and later-formed wood will therefore have higher Mesomorphy or Conductance values. The Mesomorphy index has been criticized by OEVER et al. (1981) chiefly on the grounds that it is not predictive. Certainly in the numerous papers where the Meso morphy index has been used, values are remarkably closely correlated with habitat ecology where leaves of a plant are drought-deciduous or potentially so rather than evergreen, and where no special foliar apparatus for niitigating transpiration is present. Where evergreen leaves of probable low t.ranspirational capacity (e.g., Buxus) occur, or where succulence, C4 photosynthesis, or a variety of other mecha nisms besides wood anatomy for controlling water relations are present, Mesomorphy values do not parallel the nature of the habitat. it is true. Precisely because of this, the Mesomorphy values are, in fact, predictive: they predict some device (succulence, microphylly, etc.) which mediates the water-regulation system, thereby making wood xeromorphy of lowered selective value. The Mesomorphy values for Comhretaceae by VL1ET (1979) are quite meaningful, contrary to the comments of OEvER et al., when one takes into account not merely the habitat hut also the foliage, age of sample, etc. (VLIET’s extremes for mesomorphy values are not necessary: the means alone would have offered a clearer species-by-species comparison). OEvER et al. (1981) dismiss the idea of a Mesomorphy index. saying that “func tionally incomparable units are combined”. The value of the Mesomorphy index is that the three elements are not identical in effect: narrowness of vessels is not func tionally synonymous with vessel density, nor with vessel element length: each of these three contributions independently to water conduction and water-colunm safety characteristics. While the role of shorter vessel-element length in promoting safety has been doubted (BAAS 1976; ZIaiMERMANN 1978. OEvEu et al. 1981), the role of end-walls in localizing embolisrns within shorter portions of vessels has been observed (SLATYER 1967; SPERRY 1984), and this has led to a plausible explanation (CARLQUIST 1982b) for the way in which short vessel elements may increase safety in wood. Thus, the Mesomorphy index still seems a reasonable procedure. The conductivity figure used by OEvER et al. (1981) is an alternative possibility. Tf a conductivity formula is applied t.o Staphyleaceae (Table 1, column 15), the results suggest that some Staphyleaceae are literally 1,000 times better at conduction than others, despite the habitat similarity throughout the family. The conductivity figure must therefore be misleading. if we look at the P0ISFUILLE equation from which the. conductivity formula is derived (see Oxvxn et al. 1981), we find it. does not take into account the idea of safety; rather, it says. in effect. t.he wider the tube, the bett.er the conduction. This is true for blood vessels. For wood, however, rapidity of conduction is only one aspect of water-transport characteristics. Low figures for conductivity do not mean so much that conduction is poor but that safety is of overriding value. Either Vulnerability or Mesomorhpy values (CARLQuIST 1977) tend to parallel closely ecological dimensions (e.g., rainfall or other measures of wat.er availability), whereas

level). comparable a obtain 100 to by

0.60, dividing (R Symplocaceae for = than

0.84) (R yleaceae Staph of elements = for vessel these between correlation higher a finds

one some degree; to be expected to is length and diameter cell between correlation

A Symplocos. (1981) et al. for by reported OEvER were correlations Similar initials.

canibial

fusiform from rays) of case the in indirectly, (or directly all are derived

these structures that fact the based upon are doubtless correlations these strong: is

length element tracheary imperforate and length, element vessel height., ray between

correlation The height. ray uniseriate and height, ray multiseriate diameter, element

tracheary imperforate and diameter vessel between strong also are Correlations

length. element tracheary imperforate and length element vessel between correlations

strong may

note One sought. were St.aphyleaceae of features wood among correlations

Methods),

and Materials see also (Table 2; Coefficient. Correlation Using PEARsoN’S

Correlations Statistical

study.

further needs and 1980) arboreurn (CARLQuIST Vaccinium as such dicotyledons, other

sonic in but Staphyl.ea, in not merely rings to growth related is pattern This there.

rates conductive slow t.he of because latewood in impedance no appreciable stitute

would con bars thicker The produce. bars obstacle the by minimizing earlywood in

effectiveness conductive increased to related as interpreted he can This Staphylea.

of earlywood in than latewood in thicker-walled are plates perforation on Bars

1984). (Carlquist

evidence

circumstantial judging conduction, for extent appreciable any to tion

func

apparently, do not, ordinarily Fiber-tracheids presence. tracheid with linked are

vessels solitary which in case the of instances borderline perhaps are they genera,

three these in tracheids to similar are fiber-tracheids because however, case; any in

expected be not would grouping vessel so places, in wet grow named just genera three

The vessels. grouped of value the minimizes which system. conductive subsidiary a

offer tracheids since 1984), (Carlquist tracheids of presence to related be to theorized

been have vessels Solitary solitary. nearly more are vessels Turpinia, and Staphylea,

Euscaphis, genera the In low. quite still figures are the so dry, not. very is habitat the

Obviously, dry. is habitat the the extent genera. to two those in value be selective of

would grouping 1984), vessel (Carlquist theory a recent with in accordance therefore

elements, tracheary for imperforate capability conductive poorest the have genera

two These fibers. libriform has Tapiscia liorders; pit much-reduced with tracheids

Huertea fiber has 1.23. Tu.rpinia, 1.40; scia, Staphy1eaThp 1.38; Huertea, 1.07; phis,

Eusca

follows: as by genus, are, group veselper of number for Values ficant.

signi

be may vessel) solitary a. 1 (where group per vessels 1.20 above values that

say one

may In general, vessels. of grouping minimal than more show genera two ever,

How Staphyleaceae. in degree any appreciable occur to not does grouping Vessel

1982b). (CARLQuIsT in Ilticium are they as occurs,

frost areas where to but rather shrubs, desert in are they as conditions to dry linked

to he not appear do yleaceae Staph in thickenings helical of Occurrence Staphylea.

Euscaphis and occur, they where genera in t.wo the earlywood in conspicuous least

are

thickenings helical Indeed, rings. of growth occurrence parallels yleacea.e Staph in

fiher-tracheids) sometimes (and vessels in thickenings of helical occurrence The

dimension. this

not have does equation the P0IsEuILLE increased; is availability water as capability

of conductive in favor is relinquished safety wood which way the in of representation

accurate a very seems index Mesomorphy The relations). water to related indirectly

only is and latitude used, is availability water than rather latitude that in although

1981, et

15; a!. OEvER column 1, (Table widely more much range figures conductivity

A. HOERMAN D. and CARLQuLsr Sx. 212 Wood Anatomy of Staphyleaceae 213

An even lower degree of correlation occurs between vessel element length and num ber of bars per perforation plate in both iStaphyleaceae and Symplocos. One might not have expected this, but evidently the reason for lack of correlation is that in parti cular phylads, evolutionary loss of bars is accelerated, apparently owing to selection for rapid conduction characteristics. Thus, Huertec has fewer bars than one might expect on the basis of its vessel element length, hut evidentiy has a slightly more advanced perforation plate morphology just as its fiber-tracheids have more reduced borders. This is true, in a heightened form, of Tapiscia sinensis also. A stronger correlation (an inverse one) occurs between vessel diameter and number of vessels per mm2 than between vessel diameter and vessel element length (slightly more so in Staphyleaceae, niarkedly more so in yrnplocos). This is interesting, because vessel diameter and vessel density are related to each other only by packing conside rations, which have considerable latitude, and not by morphology. However, vessel diameter is governed by degree of enlargement, whereas vessel element length is governed by length of fusiform cambial initials. As growth rings show, degree of lateral widening in vessels can vary greatly while vessel element length stays constant, or nearly so. Phylogenetic and Systematic Conclusions taphyleaceae appear to be a coherent family. The list of wood features which is common to the family is given in the Summary. While the family may be primitive in such features as relatively great length of vessel elements and large number of bars per perforation plate, &aphyleaceae can be regarded as specialized in a nunber of features. The imperforate tracheary elements represent various degrees of departure from a primitive tracheid form, depending on the genus. The ratio between vessel element length and imperforate tracheary element length is above 1.20, the figure one expects, approximately, for wood of primitive dicotyledons (CARLQUIST 1975). The rays are not of type Heterogeneous I, but Heterogeneous hA. Axial parenchvma is not diffuse (diffuse parenchyma is vestigially present), hut of various paratracheal types (ahaxial, vasicentric scanty) or ray-adjacent. The genera of Staphyleaceae can be differentiated from each other by means of wood anatomy, as summarized below. One can say that Huertea and Tapiscia are more specialized than the other three genera in having fewer bars per perforation plates and in having some (Huertea) or complete (Tapiscia) loss of borders on pits of imperforate tracheary elements. Absence of axial parenchyma in these two genera is probably a specialization, and is a.ssociated with the septate nature of iniperforate t.racheary elements. Eascaphis has growth rings, vessels with numerous (mean 31) bars per perfora tion plate, helical sculpture in latewood vessels (sometimes in earlywood vessels and fiber-tracheids as well). Imperforate tracheary elements are all fiber-tracheids with pits fully bordered but sparser than those of tracheids. Fiber-tracheids are nonseptate. Axial parenchyma is present. Multiseriate ray width is the greatest in the family. Tyloses and crystals are lacking.

Haertea lacks growth rings. Vessels are notably wide (mean = 118am) with fewer and thicker bars (mean = 14.3 bars) than the genera other than Tapiscia. and vessels Jack helical sculpture. Fiher-tracheids have much-reduced pit borders (pit cavities 2—3 m in diameter) and are septate. Axial parenchyma absent. Multiseriate rays are narrow. Tyloses are present. Crystals are present in subdivided erect ray cells and in a few chambered fiber-tracheids. S’taphylea has rather sharply demarcated growth rings. Vessel diameter (mean 49.8,um) is the smallest in the faniily, but number of bars per perforation plate is intermediate in the famil . Helical sculpture is present in vessels, especially late-

do as 1950), & CuAL1 (METCALFE parenchyma abaxial possesses (Hydrangeaceae)

Philadeiphus Tapiscia. Huertea and do as parenchyma axial lack respectively, ceae,

Grossularia and Hyclrangeaceae 1970) of al. RiSes et (STER and 1979) & S’rEmc (STYER

Deutzia yleaceac. Staph of wood of sections tangential resembles closely PLANcH.

btumei Weinmannia of section a of tangential photomicrograph (1977) DJcK1S0NS

of portions rays. in the central predominate cells proc’unihent and Staphyleacae.

those of resemble Gvnoniaceae of rays multiseriate on The 1950). wings CHALK &

Rydrangeaceae in he (METcALFE found may rays wider hut Staphyleaceae. of those

as as wide quite not usually are (uAcniaceae of Rays yleaceae. Staph in as is it vessels

around grouped (‘unoniaceae than in rather diffuse often is parenchvma Axial

1980). 1977. (D1CKIS0N fiher-t.racheids as qualify therefore which elements,

t.racheary on iniperforate pits bordered have also C’uaoniaceae of genera most and

of (‘unoniaceae.

t.hose like are Staphyleaceae of vessels primitive The 1980). DicKisoN

(see authors by various (Saxifragales) Cunoniales in placed been has Staphyleaceas

groups. smaller to than

Sapindales rather as such group polymorphic a large to family small a particular

relate to he tendency a there may so species. and genera a few only containing groups

than in is it greater understandably is diversity families, and genera sapindalean of

assetimblages large In homocellular. are rays which wide has .4ceraceae nacardiaceae. A

found he in however, may, heterocellular are rays which Wider 1950). & CHALIC

cells (METcALFE of procumbent wholly composed and wide) cells 2—3 than more

(rarely narrow are Sapindàceae Rays iii borders. no or elements tracheary perforate

on borders im vestigial the only and most on vessels plates perforation simple have

exaiiiple. for Sapindaceas. evident. strongly is specialization of degree in the disparity

Sabiaceae, than other Sapindales of that yleacsa t.o Staph of wood compares one If

relationship. phyletic to than

level rather evolutionary in similarity to in part least at he due may similarities the

but similarities, numerous the considering differences, major not are These 1950).

CHALK & crystals and (METCALFE lack rays, iltiseriate of in portions multeriseriate

in cells have erect often pitting. vessel-vessel only alternate have former the that.

yieaceae in Staph of that from differs Sabiaceae of anatomy Wood Sabiaceae. is fibers

libriform than rather and fiber-tracheids elements on vessel plates perforation form

scalari has which in order the family other only The anatomy. of wood on account

Sapindales in element discordant a are yleaceae Staph relationships, to respect With

species. one in cells ray erect vided

subdi in observed were Crystals species. in some present are Tyloses i,430,um).

(uiean family the in the tallest are mult.iseriat.es hut family the for intermediate is

width Mtiltiseriate present. ray is parenchyma Axial distributed. more sparsely ht

of as tracheid those as large pits bordered fully with fiher-tracheids nonseptate are

elements tracheary iniperforat.e All absent. is sculpture Helical genera. other the than

35.0) (mean plate

perforation per bars more with l,390ini), (mean family the =

longest in the wide, relatively are elements Vessel rings. growth lacks Turpinia

are absent.

Crystals present. are loses T family. the in 321iii) (mean =

shortest and t.he narrow relatively are rays Multiseriate absent. is parenchvma Axial

long. pits simple with 2 — 3 u1n fibers libriform septate are elements tracheary forate

All imper on vessels. absent is sculpture Helical genera. other the than 7.4) (mean

plate perforation per

bars fewer with genera. five the 4 um) of 692 mean = shortest

the bitt diameter in moderate are elements Vessel rings. growth lacks Tapi.scia

absent.

are Crystals present.. are Tyloses present. is parenchvma Axial on tracheids. those as

disposed densely as almost. pits fiillyhordered large with fiher-tracheids nonsept.ate

are all elements tracheary Iniperforate fiher-tracheids. few a in and vessels, wood

A. B. and H0EKMAN CARLQUIsP Sir. 214 ood Anatomy of Stuphyicoceae 215

Euscaphis, Staphylea. and Turpinia. Amorphous deposits are common in Cunoniaceae. Crystals occur in Hydrangeaceae, although not frequently, in wood. Thus facts of wood anatomy would not contradict a possible relationship between Staphyieaceae and Cunoniales (Saxifragales). However, both Sapindales and Cunoniales may he placed in the same superorder (TH0RNE 1981). so the question of placement of S’taphyieaceae in one of these orders versus another may not be as contrasted as it seems at first glance. References

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U.S.A. 91711 California Claremont, College,

Pomona

of Biology, Department H0EicMAN, A. DAvrn Mr. U.S.A.; 91711 California Claremont,

Garden, Botanic Asia Santa Rancho SHERWIN CARLQuIsT, Dr. Prof. addresses: Authors’

1984 11, November Received

Cambridge.

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