~ ( 

~ 

i! 

'I 

COMPARATIVE WOOD ANATOMY OF 

SOME SHRUBS NATIVE TO THE 

'NORTHERN ROCKY MOUNTAINS 

ARLENE DALE 

RESEARCH PAPER INT-45, 1968 

Intermountain Forest and Range 

Experiment Station 

Ogden, Utah 

This file was created by scanning the printed publication.

Errors identified by the software have been corrected;

however, some errors may remain.
THE AUTHOR 

MRS. ARLENE DALE has been a research technician in the 

Department of Botany, University of Montana, Missoula, 

since 1964. Most of her work deals with histological 

studies of fungi- -primarily the needlecast of conifers 

--and of plants damaged by air pollutants. 

This paper is based on content of the thesis submitted by 

the author as partial fulfillment of the requirements for 

the Master of Arts degree in Botany at the University of 

Montana. This project was supported jointly by the 

University of Montana and the U.S. Forest Service, 

Intermountain Forest and Range Experiment Station. 
COMPARATIVE WOOD 

U.S. Forest Service 

Research Paper INT -45 

1968 

ANATOMY 

OF SOME SHRUBS 

NATIVE TO 

THE NORTHERN ROCKY MOUNTAINS 

ARLENE DALE 

INTERMOUNTAIN FOREST AND RANGE EXPERIMENT STATION 

Forest Service 

U.S. Department of Agriculture 

Ogden, Utah 84401 

Joseph F. Pechanec, Director 
INTRODUCTION . • . • • . . 

METHODS AND MATERIALS 

Field Collections . • . . 

CONTENTS 

Specimen Preparation • 0 • 

Preparation of Key • . 

Discussion . . . • • • • . 0 • • 

GENERA INCLUDED IN KEY BY FAMILY 

KEY TO SOME SHRUB WOODS NATIVE TO THE 

NORTHERN ROCKY MOUNTAINS .. 

PHOTOMICROGRAPHS AND SYNOPSES OF CHARACTERS 

USED IN KEY .•..•. 

DIAGRAMS OF WOOD CHARACTERS . • o • 

GLOSSARY • . . . ..•...•. 

REFERENCES. • . o • • • • • o • • 

SPECIES INDEX • o • • • o • • • 

Page 

1 

1 

1 

2 

2 

2 

3 

4 

7 

66 

69 

72 

73 
. ,1;· __ 

COMPARATIVE WOOD ANATOMY OF SOME SHRUBS NATIVE TO 

THE NORTHERN ROCKY MOUNTAINS 

Arlene Dale 

INTRODUCTION 

Presented here is a key for identification of shrubby plants native to the Northern Rocky 

Mountain area. This key supplements those already in existence that are based on plant organs, 

such as flowers, leaves, fruits, and other macroscopic characters. In contrast, the characters 

upon which this key is based are those of the wood, or technically, the xylem. Such a key would 

be valuable to substantiate identifications made by means of the ordinary keys and to aid in the 

identifications of plants when flowers, leaves, and fruits are not available . 

Fifty-five species, representing 35 genera (see page 3), have been incorporated in this 

key. Xylem characters determined the arrangement of the species. Where certain characters 

were held in common by all genera of a family, these species parallel their usual taxonomic 

arrangement. Depending on characters present, identification is to a species, a genus, or a 

group of genera. Further study will make possible more detailed identification. 

A review of the literature concerning wood anatomy indicates a change in emphasis has 

occurred within the past 20 years- -from a heavy reliance on quantitative data in distinguishing 

among woods to the use of qualitative characters. This change has been caused by increased 

understanding of the extent and causes of variations within a species. No longer is it assumed 

that a single specimen accurately represents a species. To determine the range of variability 

of all 55 species was beyond the scope of this study; the diversity of characters within each 

species will be discovered only as this key is used. In this key, cell patterns-- created by the 

ordered activity of the cambium--are employed extensively. 

METHODS AND MATERIALS 

FIELD COLLECTIONS 

Specimens wer:_e collected for over 60 species of shrubs from sites in western Montana 

and northern Idaho.-'- Several of these species were rejected from consideration because they 

were represented by only one specimen; others were rejected because the specimens collected 

were only 1 year old. While some species were represented by as many as five specimens, the 

minimum number was two. 

1 Most of the collections and all of the identifications were made by Peter F. Stickney, 

Associate Plant Ecologist, U.S. Forest Service, Intermountain Forest and Range Experiment 

Station. Identifications were made by means of the following: Hitchcock, C. L., A. Cronquist, 

M. Ownbey, and J. W. Thompson. Vascular plants of the Pacific Northwest. Seattle: Univ. 

Wash. Press. Part 2: Salicaceae to Saxifragaceae, 1964; Part 3: Saxifragaceae to Ericaceae, 

614 pp., 1961; Part 4: Ericaceae through Campanulaceae, 510 pp., 1959; Part 5: Compositae, 

314 pp . ' 1 9 55 . 
SPECIMEN PREPARATION 

The paraffin -embedding method was employed in preparing the microscopic slides from 

the specimens. Small wedges were cut from the specimens, aspirated, dehydrated through a 

tertiary butyl alcohol series, and embedded in Paraplast.2 After soaking in an alcohol- glycerine 

solution of Aerosol OT 100, the embedded specimens were sectioned in thicknesses of from six 

to 20 micra on a rotary microtome; for most of the material a thickness of eight to 10 micra 

was optimum. The paraffin ribbons were affixed to slides with Haupt's adhesive. After removal of the paraffin by xylene, the sections were run through an ethyl-alcohol series into 

water, stained by means of the Feulgen reaction, dehydrated in a second alcohol series, and 

counterstained with fast green. In the first alcohol series, a celloidon solution was interposed 

to improve adhesion of sections. The mounting medium was Harleco 's Synthetic Resin. Cross, 

radial, and tangential sections were cut for each species. 

PREPARATION OF KEY 

The key was first outlined from photomicrographs of the sections and then checked with 

the slides and with published accounts of wood anatomy. Because in many cases the original 

photographs did not show the characters sought, the slides for each species had to be examined 

several times. Each character used to distinguish a species was checked in all specimens 

representing that species. 

The characters used to distinguish among the species are not in all cases the same; vessel distribution, perforation plates, rays, and axial parenchyma are the characters most often 

utilized. Where information about wood anatomy for a species was available, it was consulted 

as a check against the specimens used in this study. These accounts, with few exceptions, 

confirmed the conclusions outlined. 

DISCUSSION 

The premise upon which this study was based is that species of shrubs are identifiable on 

the basis of characteristics of the xylem. However, evidence indicates that such distinction 

cannot be made. The results of this study confirm that in many genera the species produce 

xylem that is so similar as to be valueless for identification; e.g., the two species of Ceanothus 

included in this key. Consequently, wood traits alone may not be adequate to identify a shrub. 

Before attempting to use this key, the reader should recognize that identification of shrubs 

by means of microscopic characteristics of the wood is difficult. Careful preparation of specimens and adequate laboratory facilities are necessary. Moreover, error may result from two 

sources. The first is that the unknown shrub may be one not included in this study. The second 

is that spec_ies variability has not been assessed. Identification of unknown shrubs by xylem 

should be attempted only when all other means have been exhausted. 

2 Use of trade names herein is for identification only, and does not imply endorsement by 

the U.S. Forest Service. 

2 
GENERA INCLUDED IN KEY BY FAMILY 3 

Taxaceae 

Tax us 

Pinaceae 

Juniperus 

Compo sitae 

Artemisia 

Chrysothamnus 

Rosaceae 

Amelanchier 

Cercocarpus 

Crataegus 

Holodiscus 

Physocarpus 

Prunus 

Purshia 

Rosa 

Rubus 

Sorbus 

Spiraea 

Saxifragaceae 

Philadelphus 

Ribes 

Aceraceae 

Acer 

Salicaceae 

Populus 

Salix 

Betulaceae 

Alnus 

Betula 

Cornaceae 

Corn us 

Ericaceae 

Arctostaphylos 

Ledum 

Menziesia 

Vaccinium 

Anacardiaceae 

Rhus 

Rhamnaceae 

Ceanothus 

Rhamnus 

Caprifoliaceae 

Lonicera 

Sambucus 

Symphoricarpos 

Elaeagnaceae 

Shepherdia 

Celastraceae 

Pachistima 

3 As classified in Lawrence, George H. M. Taxonomy of 

Vascular Plants. New York: The MacMillan Co. 1951. 

3 
KEY TO SOME SHRUB WOODS NATIVE TO THE NORTHERN ROCKY MOUNTAINS 

1 Vessels present • • • • • • • o • o • • • • • • • o • • • • • o • • • • • • 3 

1 Vessels not present • • o • • o • • • • • • • • • o o • • • • o • • 2 

2 Tracheids with spiral thickenings • • • • • • Taxus brevifolia (p. 8- 9) 

2 Tracheids without spiral thickenings. . jiilliperus spp. (p. 8-9) 

3 Rays 3 to many cells in width, often massive, dispersed between uniseriate rays. 4 

3 Rays 1 or 2 cells in width (rarely 3) • • • • • • • • • • • • • • • • • • • • • . • 17 

4 Vessels grouped in patterns forming radial or concentric bands or arcs . . • • . 5 

4 Vessels predominantly solitary or in pairs, except for initial vessels of growing 

season which may be contiguous • • . . • • . • • • • • • • • • • • • . . 8 

5 Vessel bands radial • o ••• o ••• o o • • • Prunus spp. (p. 16-17, 18-19) 

5 Vessel bands concentric or vessels forming arcs • • • • • • • . . o • • • • • 6 

6 Band of cork tissue initiating each growth ring Artemisia tridentata (p. 8- 9) 

6 Band of cork not present as above . . . • • • • • • . • . • • . . . • . . 7 

7 Vessels in concentric bands, alternating with bands of 

tracheids and fibers • • o • • • • o • • • 0 • • • • Chrysothamnus nauseosus (p. 10) 

7 Vessels in slight arcs rather than in concentric bands . Chrysothamnus viscidiflorus 

(p. 10-11) 

8 Perforation plates simple • • . • • • . • • • • . . • • • . . . • • . . • 9 

8 Perforation plates scalariform . • • . . 0 • • • • • • • • • • • • • • • • • 11 

9 Growth ring distinct; vessels decreasing in size from early to late wood; vessels 

relatively numerous • • • • • • . o • • • o • • • • • • • • • • • • • • • • 10 

9 Growth ring indistinct; vessels not changing in size or spacing from early to late 

wood; vessels relatively few and far apart . • Holodiscus discolor (p. 14-15) 

or Rubus spp. (p. 22-23, 24-25) 

10 Vessels markedly larger in early than in 

late wood • . o • • • • • o • • • • o • Rosa woodsii (p. 20-21) 

10 Vessels not markedly larger in early than 

in late wood . • • • • . • . • • • . • • o Rosa gymnocarpa (p. 20-21) 

11 Tracheids and fibers alternating in radial bands Philadelphus lewisii (p. 26- 27) 

11 Tracheids not alternating with fibers as above • • • • o • • • • • • o • o 12 

13 

13 

15 

15 

4 

12 Vessels forming complete or short concentric bands within growth ring, 

usually 1 vessel in width • • . . • • . • o • • • • • • • • • • • • o • • • • 13 

12 Vessels not forming such bands, instead being more in diagonal bands 

or diffuse • . . • • • • • . . o o 0 • • • o 

Bands complete; vessels of successive bands in any 

one growth ring are of similar size • • 0 • • • • 

Bands tending to be interrupted not forming complete 

bands; vessels of successive bands in any one growth 

14 

Ribes aureum (p. 28-29) 

not similar in size; the initial vessels usually are . 

markedly larger than succeeding ones • • • • • • Ribes cereum (p. 28- 29) 

14 Ve-ssels mostly solitary with some small clusters; are randomly dispersed, 

with no clear pattern evident . • • • • • • • • • • • • • • • • • • • . • . • 15 

14 Vessels arranged in diagonal bands or arcs with a few short, tangential bands . • 16 

Growth ring initiated with a band of larger vessels; the 

vessels become progressively smaller through the 

growing season; vessels relatively few. • . • • . . • Ribes lacustre (p. 32-33, 34-35) 

Growth ring not initiated with a band of larger vessels 

although the vessels of the last-formed wood are noticeably smaller than the initial vessels, making the growth 

rings discernible • . • . • o • • 0 • o • • • • • o Ribes hudsonianum (p. 30-31) 
17 

17 

19 

19 

21 

21 

23 

23 

25 

25 

27 

27 

28 

28 

29 

29 

31 

16 Vessels forming a zigzag pattern, radially alined . Ribes viscosissimum (p. 34-35) 

16 Vessels may be diagonally alined, in short 

tangential bands, and/or diffuse, with none of 

these found exclusively; there is not the regularity 

of alternating diagonal bands as in the above • • • • Ribes irriguum (p. 30-31, 34-35) 

or Ribes setosum (p. 32 -33) 

Vessels commonly in radial multiples; often spring 

wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single 

vessel), the vessels progressing toward formation of 

radial multiples from early to late wood. . • • o • o • • • • • • • • • • • • 

Vessels solitary, in pairs, or in various aggregates, including sometimes radial 

18 

vessel chains but not vessel multiples • • . o • • • • • • • 24 

18 Perforation plates simple . . . • • . . • • 0 • 

18 Perforation plates scalariform . • • • . • • • • . • • . . • . • • • • 

Rays composed of procumbent cells only . 0 • • 

Rays composed of both procumbent and upright cells 

20 Vessels with spiral thickenings; rays mainly 

uniseriate but some 2 to 4 cells wide . • 

20 Vessels without spiral thickenings; rays 

. Acer glabrum (p. 36-37) 

0 19 

• 22 

• 20 

• 21 

uniseriate only o • o o • Populus spp. (p. 38-39, 40-41) 

Rays 1 cell in width o • • • • o • • • Salix scouleriana (p. 40-41) 

Rays 2 to 4 cells in width . . . . . ... 0 • Sambucus spp. (p. 60-61, 62-63) 

22 Rays uniseriate only. . . 0 • • • • • • • • • • • • • 0 • • 23 

22 Rays both 1 and 2 cells in width .......... Betula spp. (p. 44-45) 

Axial parenchyma forming tangential bands at short 

intervals between rays .......... 0 ••••• Alnus incana (Po 42-43) 

Axial parenchyma sparse and diffuse . . . . . . Alnus sinuata (p. 42-43, 46-4 7) 

24 Vessel perforation plates scalariform 0 • • 25 

24 Vessel perforation plates simple . . . . 0 • • • • • • • • • • • 28 

Vessels regular in size and spacing throughout 

growth ring ..................... Cornus stolonifera (p. 46-47) 

Vessels not regular in size or spacing as above . 0 • • • • • • • • • 26 

26 Rays 1 to several cells wide ....... 0 • 0 • Vaccinium spp. (p. 50-51, 52-53) 

26 Rays uniseriate only. . 0 • • • • • • • • • • • • • • • • • • • • • • • • • 27 

Growth ring initiated by a single row of large, 

closely spaced vessels •....... 0 •••••• 0 Menziesia ferruginea (p. 50-51) 

Growth ring not initiated as above; no clear line 

of large vessels. . . . . o • • • • • • • • • • 0 • • Ledum glandulosa (p. 48-49) 

Annual ring initiated by a definite band of large vessels; other vessels are much 

smalle_r and sparse, giving a two-banded appearance to the growth ring. . . . . 

Annual ring not initiated by band of large vessels as above; vessels grade into 

smaller sizes as the growing season progresses without an abrupt change in size 

29 

30 

Axial parenchyma numerous, paratracheal, and diffuse, 

often encircling the larger vessels • . . . . . 0 Symphoricarpos albus (p. 62-63) 

Axial parenchyma few, if in contact with a vessel, a 

single cell rather than an encircling band . 0 • • • Arctostaphylos uva -ursi (p. 48-49) 

30 Axial parenchyma predominantly paratracheal 0 . • . . . 31 

30 Axial parenchyma predominantly terminal or diffuse 0 • • • • • • • • • • • • 32 

Axial parenchyma distributed as 2 to several cells on the 

periphery of each vessel group; vessels solitary or in 

5 
clusters of a few cells in the early wood progressing to 

radial files which are grouped secondarily to form tangential bands in the late wood . • • • • • . • . . . • Rhus glabra (p. 52- 53) 

31 Axial parenchyma distributed as an occasional single 

33 

33 

cell adjacent to a vessel; vessels diffuse, not grouped 

as above • • • • . . • • . o • • • • • • • • 0 • • 

32 Axial parenchyma predominantly terminal . . • 

32 Axial parenchyma predominantly diffuse or absent 

Vessels decreasing gradually in size from early to late 

wood; vessels smooth in outline, solitary, or in occasional 

Pachistima myrsinites (p. 64 -65} 

• • • • 0 • • • 33 

35 

pairs • • 0 • • • • • • • o • • • • 0 • • • • • • • Physocarpus malvaceus (p. 16 -17) 

Vessels not decreasing gradually in size as above; vessels angular in outline. . • • . 34 

34 Vessels arranged in diagonal and/or radial lines . Rhamnus alnifolia (p. 56- 57) 

34 Vessels solitary or in occasional multiples of a 

few cells . • . o • • • • • • • • • • • • • • • Rhamnus purshiana (p. 56-57) 

35 Vessels occurring in patterns of radially alined S curves 

in mature wood; tannins occurring in even the young ray 

parenchyma •••. o o ••• o •••••• o o • • Ceanothus spp. (p. 54-55) 

35 Vessels not in patterns as described above; tannins, if present, occurring in ray 

parenchyma only in the more mature cells . . 36 

36 Axial parenchyma numerous . • . 0 o 0 • • • • • • • • • • • 0 0 • • • 37 

36 Axial parenchyma absent or sparse • • • • • • • • . • . • . . • • . 0 • 39 

37 Vessel walls composed of five or six facets, giving the vessel a definitely angular 

37 

39 

39 

'41 

41 

43 

43 

6 

appearance • • . o o o o • • o • • • • • • • • • • • • • • • • • • • • 0 • 38 

Vessel walls, although composed of several facets, giving 

a smoother, more rounded outline than above. . . . • Amelanchier alnifolia (p. 12 -13) 

38 Vessels tending to form concentric bands within the 

growth ring; vessels relatively few and large. • • Shepherdia canadensis (p. 64-65) 

38 Vessels diffuse in distribution with no particular 

alinement; vessels relatively small and numerous. Crataegus douglasii (p. 14-15) 

Spring wood vessels large, numerous, and solitary 

(although closely spaced); vessels of late wood not 

greatly smaller than those of early wood, but much 

more widely spaced; vessels markedly angular, with 

concave facets .•••.••••.•..•. o o • Sorbus scopulina (Po 24-25) 

Spring wood vessels not as above; if vessel spacing is as above, then vessels are 

rounded, with convex facets; if vessel shape is as above, then spacing differs 

40 Early wood with a few large vessels; vessel size 

decreases greatly from early to late wood, but 

40 

spacing of vessels remains similar . • . . . Cercocarpus ledifolius (p. 12 -13) 

40 Early wood may have a few large vessels but, if so, then the vessel size and 

spacing differ from the above • • • . • • • • 41 

Axial parenchyma absent o o o • • • • 0 • • • • Spiraea betulifolia (p. 26-27) 

Axial parenchyma present although few in number. 0 • • • • • 0 • • • • • 0 42 

42 Vessels angular in outline . • . 0 • • 0 

42 Vessels rounded in outline o • • 0 • • • 

Ray cells squared in outline in cross section . 

Ray cells radially elongated in cross sections 

43 

Purshia tridentata (p. 18 -19) 

Lonicera utahensis (p. 58-59) 

Lonicera involucrata (p. 58-59) 
PHOTOMICROGRAPHS 

AND 

SYNOPSES OF CHARACTERS 

USED IN KEY 

ALL PHOTOMICROGRAPHS ARE OF CROSS 

SECTIONS UNLESS OTHERWISE NOTED AS: 

* radial 

* * tangential 

7 
8 

PLATE I 

Taxus brevifolia (la and lb*) 

Vessels absent; tracheids with spiral thickenings. 

Juniperus communis (2a) and 

Juniperus scopulorum (3a) 

Vessels absent; tracheids without spiral thickenings. 

Artemisia tridentata (4a and 4b) 

Vessels present, grouped in patterns forming concentric 

bands; rays three to many cells in width, often massive, 

dispersed between uniseriate rays; band of cork tissue 

initiating each growth ring. 
9 
10 

PLATE II 

Chrysothamnus nauseosus (6a and 6b} 

Vessels present, grouped in patterns forming concentric 

bands; rays three to many cells in width, often massive, 

dispersed between uniseriate rays. 

Chrysothamnus viscidiflorus (Sa and Sb} 

Vessels present, grouped in patterns forming slight arcs; 

rays three to many cells in width, often massive, dispersed 

between uniseriate rays. 
11 
12 

PLATE III 

Amelanchier alnifolia (7 a, 7b, and 7 c *) 

Vessels present, solitary or in pairs; annual ring not initiated 

by band of large vessels; vessels grade into smaller sizes as 

the growing season progresses without an abrupt change in 

size; vessel walls giving a smooth, rounded outline to vessel 

in cross section; vessel perforation plate simple; axial parenchyma numerous, diffuse; rays one to two cells in width (rarely 

three). 

Cercocarpus ledifolius (8a and 8b) 

Vessels present, solitary, in pairs or various aggregates; 

annual ring not initiated by a band of large vessels; vessels 

grade into smaller sizes as the growing season progresses 

without an abrupt change in size; early wood with a few large 

vessels; vessel size decreases greatly from early to late wood, 

but spacing of vessels remains similar; perforation plate simpie; axial parenchyma absent or sparse and diffuse; rays one 

or two cells in width (rarely three). 
' .- . ~ ... '' 

13 
14 

PLATE IV 

Crataegus douglasii (9a and 9b) 

Vessels present, diffuse in distribution with no particular 

alinement; vessels relatively small and numerous; vessel 

walls composed of five or six facets, giving the vessel a 

definitely angular appearance; vessels grade into smaller 

sizes as the growing season progresses without an abrupt 

change in size; vessels solitary or in pairs, or in various 

aggregates; vessel perforation plates simple; axial parenchyma absent or sparse, diffuse; rays one or two cells in 

width (rarely three). 

Holodiscus discolor (lOa and lOb) 

Vessels present; vessels predominantly solitary or in pairs, 

except for initial vessels which may be contiguous; growth 

ring indistinct; vessels not changing in size or spacing from 

early to late wood; vessels relatively few and far apart; perforation plates simple; rays three to many cells in width, 

often massive, dispersed between uniseriate rays. 
lS 
16 

PLATE V 

Physocarpus malvaceus (lla and llb) 

Vessels present; vessels decreasing gradually in size from 

early to late wood; vessels smooth in outline, solitary or in 

occasional pairs; vessel perforation plates simple; axial 

parenchyma predominantly diffuse or absent; rays one or 

two cells in width (rarely three). 

Prunus emarginata (12a, 12b, and 12c*) 

Vessels present; vessels grouped in radial bands; rays 

three to many cells in width, dispersed between uniseriate 

rays. 
17 
18 

PLATE VI 

Prunus virginiana (13a and 13b) 

(Same synopsis as for Prunus emarginata, page 16) 

Purshia tridentata (14a and 14b) 

Vessels present; vessels rounded in outline; vessels grade 

into smaller sizes as the growing season progresses without an abrupt change in size; vessels solitary, in pairs, or 

in various aggregates; perforation plates simple; axial 

parenchyma present although few in number, diffuse; rays 

one or two cells in width (rarely three). 
19 
20 

PLATE VII 

Rosa gymnocarpa (15a and 15b) 

Vessels present; vessels not markedly larger in early than 

in late wood; growth ring distinct; vessels decreasing in 

size from early to late wood; vessels relatively numerous; 

vessels predominantly solitary or in pairs, except for initial 

vessels which may be contiguous; perforation plates simple; 

rays three to many cells in width, dispersed between uniseriate rays. 

Rosa woodsii (16a and 16b) 

Vessels present; vessels markedly larger in early than in 

late wood; growth ring distinct; vessels decreasing in size 

from early to late wood; vessels relatively numerous; vessels predominantly solitary or in pairs, except for initial 

vessels which may be contiguous; perforation plates simple; 

rays three to many cells in width, dispersed between uniseriate rays. 
2! 
22 

PLATE VIII 

Rubus idaeus (17a, 17b, and 17c**) and 

Rubus leucoderrnjs (18a and 18b) 

Vessels present; vessels predominantly solitary or in pairs, 

except for initial vessels which may be contiguous; growth 

ring indistinct; vessels not changing in size or spacing from 

early to late wood; vessels relatively few and far apart; perforation plates simple; rays three to many cells in width, 

dispersed between uniseriate rays. 
23 
24 

PLATE IX 

Rubus parviflorus (19a and 19b) 

(Same synopsis as for Rubus leucodermis, page 22) 

Sorbus scopulina (20a and 20b) 

Vessels present; spring wood vessels large, numerous and 

solitary (although closely spaced); vessels of late wood not 

much smaller than those of early wood, but much more 

widely spaced; vessels markedly angular, with concave 

facets; perforation plate simple; axial parenchyma absent 

or sparse and diffuse; rays one or two cells in width (rarely 

three). 
25 
'· .... 

26 

PLATE X 

Spiraea betulifolia (2la and 2lb) 

Vessels present; vessels grade into smaller sizes as the 

growing season progresses; without an abrupt change in 

size; vessels solitary, in pairs, or in various aggregates; 

perforation plates simple; axial parenchyma absent; rays 

one or two cells in width (rarely three). 

Philadelphus lewisii (22a and 22b) 

Vessels present; vessels predominantly solitary or in pairs, 

except for initial vessels which may be contiguous; perforation plates scalariform; rays three to many cells in width, 

often massive, dispersed between uniseriate rays; tracheids 

and fibers alternating in radial bands. 

28 

PLATE XI 

Ribes aureum (23a and 23b) 

Vessels present; vessels forming complete concentric bands 

within the growth ring, usually one vessel in width, vessels 

of successive bands in any one growth ring are of similar 

size; vessels predominantly solitary or in pairs, except for 

initial vessels which may be contiguous; perforation plates 

scalariform; rays three to many cells in width, often massive, dispersed between uniseriate rays. 

Ribes cereum (24a and 24b) 

Vessels present; vessels forming interrupted concentric 

bands within the growth ring, usually 1 vessel in width; 

vessels of successive bands in any one growth ring not 

similar in size; the initial vessels are usually markedly 

larger than succeeding ones; vessels predominantly solitary or in pairs, except for initial vessels which may be 

contiguous; perforation plates scalariform; rays three to 

many cells in width; often massive, dispersed betwwen 

uniseriate rays. 
29 
··;,;_:, 

30 

PLATE XII 

Ribes hudsonianum (2 Sa and 2 Sb) 

Vessels present; vessels mostly solitary with some small 

clusters of vessels; these vessels or clusters are randomly 

dispersed with no clear pattern evident; growth ring not initiated with a band of larger vessels although the vessels of 

the last-formed wood are noticeably smaller than the initial 

vessels, making the growth rings discernible; perforation 

plates scalariform; rays three to many cells in width, often 

massive, dispersed between uniseriate rays. 

Ribes irriguum (26a and 26b) 

Vessels present; vessels may be diagonally alined, in 

short tangential bands, and/or diffuse, with none of these 

found exclusively; perforation plate scalariform; rays three 

to many cells in width, often massive, dispersed between 

uniseriate rays. 
I/ 

-

.; -· 

V : I 

31 
32 

PLATE XIII 

Ribes lacustre (27a and 27b) 

Vessels present; growth ring initiated with a band of larger 

vessels; the vessels become progressively smaller through 

the growing season; vessels relatively few; vessels predominantly solitary, in pairs, or in various aggregates; perforation plate scalariform; rays three to many cells in width, 

often massive, dispersed between uniseriate rays 0 

Ribes setosum (28a and 28b) 

(Same synopsis as for Ribes irriguum, page 30) 
33 
34 

PLATE XIV 

Ribes viscosissimum (29a and 29b) 

Vessels present; vessels predominantly solitary, in pairs, 

or in various aggregates; vessels forming a zigzag pattern, 

radially alined; perforation plates scalariform; rays three to 

many cells in width, often massive, dispersed between uniseriate rays. 

Ribes irriguum (26c*) 

(See page 30) 

Ribes lacustre (27c**) 

(See page 32) 
35 
36 

PLATE XV 

Acer glabrum (30a, 30b, 30c**, and 30d*) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular clusters (some appearing as subdivisions of a single vessel), the vessels progressing toward formation of radial 

multiples from early to late wood; vessels with spiral thickenings; perforation plates simple; rays composed of procumbent cells only; rays mainly uniseriate but some two to four 

cells wide. 
37 
38 

PLATE XVI 

Populus tremuloides (3la and 3lb) and 

Populus trichocarpa (32a and 32b) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular clusters (some appearing as subdivisions of a single vessel) , 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessels without spiral thickenings; 

perforation plates simple; rays composed of procumbent cells 

only; rays uniseriate only. 
39 
40 

PLATE XVII 

Salix scouleriana (33a, 33b, and 33c*) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single vessel), 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessel perforation plates simple; rays 

composed of both procumbent and upright cells; rays uniseriate 

only. 

Populus trichocarpa (32c*) 

(See page 3 8) 
,·. 

4 1 
42 

PLATE XVIII 

Alnus incana (34a and 34b) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single vessel), 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessel perforation plates scalariform; axial parenchyma forming tangential bands at short 

intervals between rays; rays uniseriate only. 

Alnus sinuata (35a and 35b) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single vessel), 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessel perforation plates scalariform; 

axial parenchyma sparse and diffuse; rays uniseriate only. 
43 
44 

PLATE XIX 

Betula glandulosa (36a and 36b) and 

Betula occidentalis (37a and 37b) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single vessel), 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessel perforation plates scalariform; 

rays both one and two cells in width. 
45 
46 

PLATE XX 

Cornus stolonifera (38a, 38b, and 38c*) 

Vessels present; vessels solitary or in pairs (appearing as a 

bisected vessel); vessels regular in size and spacing throughout growth ring; perforation plates scalariform; rays one or 

two cells in width (rarely three). 

Alnus sinuata (35c*) 

(See page 42) 

,, :··,·. 
47 
48 

PLATE XXI 

Arctostaphylos uva-ursi (39a, 39b, and 39c*) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; annual ring initiated by a definite band of large 

vessels; other vessels are much smaller and sparse, giving 

a two- banded appearance to the growth ring; perforation plates 

simple; axial parenchyma few, if in contact with a vessel, a 

single cell rather than an encircling band; rays one or two 

cells wide (rarely three). 

Ledum glandulosa (40a and 40b) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessel perforation plates scalariform; rays uniseriate only. 
49 
50 

PLATE XXII 

Menziesia ferruginea (4la, 4lb, and 4lc*) 

Vessels present; growth ring initiated by a single row of large, 

closely spaced vessels; vessels solitary, in pairs, or in various aggregates; perforation plate scalariform; rays uniseriate 

only. 

Vaccinium membranaceum (42a and 42b) 

Vessels present; vessels solitary, in pairs, or in various aggregates; perforation plate scalariform; rays one to several 

cells wide. 
51 
52 

PLATE XXIII 

Vaccinium scoparium (43a, 43b, and 43c*) 

(Same synopsis as for Vaccinium membranaceum, page 50) 

Rhus glabra ( 44a and 44b) 

Vessels present; vessels solitary, in pairs, or in various aggregates; vessels grade into smaller sizes as the growing 

season progresses without an abrupt change in size; vessels 

solitary or in clusters of a few cells in the early wood progressing to radial files which are grouped secondarily to form 

tangential bands in the late wood; perforation plate simple; 

axial parenchyma distributed as two to several cells on the 

periphery of each vessel group; rays one or two cells wide 

(rarely three). 
53 
54 

PLATE XXIV 

Ceanothus sanguineus (45a and 45b) and 

Ceanothus velutinus (46a and 46b) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessels grade into smaller sizes as the growing 

season progresses without an abrupt change in size; vessels 

occurring in patterns of radially alined S curves in mature 

wood; perforation plates simple; axial parenchyma predominantly diffuse; rays one or two cells in width (rarely three); 

tannins occurring in even the young ray cells. 

:· .. · 
ss 
56 

PLATE XXV 

Rhamnus alnifolia (47a and 47b) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessels grade into smaller sizes as the growing 

season progresses without an abrupt change in size; vessels 

arranged in diagonal and/or radial lines; vessels angular in 

outline; perforation plates simple; axial parenchyma predominantly terminal rays one or two cells in width (rarely three). 

Rhamnus purshiana (48a, 48b, and 48c*) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessels grade into smaller sizes as the growing 

season progresses without an abrupt change in size; vessels 

solitary or in occasional multiples of a few cells; vessels 

angular in outline; perforation plates simple; axial parenchyma predominantly terminal rays; one or two cells in 

width (rarely three). 
57 
58 

PLATE XXVI 

Lonicera involucrata (49a and 49b) 

Vessels present; vessels angular in outline; vessels solitary, 

in pairs, or in various aggregates; vessels grade into smaller 

sizes as the growing season progresses without an abrupt 

change in size; vessel perforation plates simple; axial parenchyma present though sparse and diffuse; rays one or two cells 

in width (rarely three), ray cells radially elongated in cross 

sections. 

Lonicera utahensis (50a, SOb, and SOc*) 

Vessels present; vessels angular in outline; vessels solitary, 

in pairs, or in various aggregates; vessels grade into smaller 

sizes as the growing season progresses without an abrupt 

change in size; vessel perforation plates simple; axial parenchyma present though sparse and diffuse; rays one or two cells 

in width (rarely three); ray cells squared in outline in cross 

sections. 
59 
60 

PLATE XXVII 

Sambucus cerulea (5la and 5lb) and 

Sambucus racemosa (52a and 52b) 

Vessels present; vessels commonly in radial multiples; often 

spring wood vessels are closely spaced singles or in irregular 

clusters (some appearing as subdivisions of a single vessel), 

the vessels progressing toward formation of radial multiples 

from early to late wood; vessel perforation plates simple; rays 

composed of both procumbent and upright cells; rays two to 

four cells in width. 
61 
62 

PLATE XXVIII 

Sambucus racemosa (52c* and 52d**) 

(See page 60) 

Symphoricarpos albus (53a and 53b) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; annual ring initiated by a definite band of large 

vessels; other vessels are much smaller and sparse, giving 

a two-banded appearance to the growth ring; perforation 

plates simple; axial parenchyma numerous, paratracheal and 

diffuse, often encircling the larger vessels, rays one or two 

cells wide (rarely three). 
63 
64 

PLATE XXIX 

Shepherdia canadensis (54a and 54b) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessels tend to form concentric bands within the 

growth ring; vessels relatively few and large; vessel walls 

composed of five or six facets, giving the vessel a definitely 

angular appearance; vessels grade into smaller sizes as the 

growing season progresses without an abrupt change in size; 

vessel perforation plates simple; axial parenchyma absent or 

sparse, diffuse; r<;1.ys one or two cells in width (rarely three). 

Pachistima myrsinites (55a and SSb) 

Vessels present; vessels solitary, in pairs, or in various 

aggregates; vessels grade into smaller sizes as the growing 

season progresses without an abrupt change in size; vessels 

diffuse; perforation plate simple; axial parenchyma distributed 

as an occasional single cell adjacent to a vessel; rays one or 

two cells wide (rarely three). 
65 
A 

D 

0 

0 C> ~ 

I 

I 

I 

I 

I 

I 

I 

I 

I 

I 

0 0 

c .. Q 0 

b 

I 

B 

b 

c 

() " 

.. a 

0 " 

Cl 0 

c 

E 

PLATE XXX. DIAGRAMS OF WOOD CHARACTERS. A, (a) procumbent ray cells in radial 

view; (b) upright ray cells in radial view. ~, (a) simple perforation plate; (b) scalariform 

perforation plate. C, (a) procumbent ray cells in tangential view; (b) upright ray cells in 

tangential view o D, (a) cross or transverse view; (b) radial view; (c) tangential view. 

E, xylem elements as seen in cross view: (a) vessel; (b) axial parenchyma; (c) ray parenchyma; (d) tracheid; (e) fiber o 

66 
B 

c 

ooo 

oo 

0 

0 a@) 0 

00 0 0 

000 a b 

D G 

0 0 0 (]J 0 

0 0 

0 

0 0~ 

H 

0 

0 0 

0 0 0 

000 

000 

E 

0 0 0 

0 0 

0 0 

K 

@ 0 0 0 ® w . 0 0 0 c:D @ 0 

0 0 

Q!} 1'9 0 

@ 0 0 

© o·· 0 . 

Q2) 00 0 

0 ® ooo 0 @ 

F I L 

PLATE XXXL DIAGRAMS OF WOOD CHARACTERS. A, axial parenchyma distribution: (a) 

diffuse; (b) paratracheal; (c) terminal. ~' axial parenchyma few. C, axial parenchyma 

numerous. D, vessels not changing in size in growth ring. .§_, vessels changing gradually in size in growth ring. F, vessels changing abruptly in size in growth ring. G, radial 

vessel alinement: (a) radial chain; (b) radial multiple. H, types of vessel pairs.-!_, solitary vessels. _L, spacing of vessels increases in growth ring. K, spacing of vessels remains similar in growth ring. L, types of vessel aggregates. 

67 

. ' .. 

GLOSSARY4 

Cell. --A chamber or compartment at some time containing a protoplast; cells form the structural units of plant tissues. 

Fibre, Fiber (Am.). --A general term of convenience in wood anatomy for any long, narrow 

cell of wood or bast other than vessels and parenchyma. Note: Often further qualified 

as wood fibres or bast fibres; the former including both the tracheids of gymnosperms 

and the libriform wood fibres and fibre -tracheids of woody angiosperms. Also used 

loosely for wood elements in general. 

Growth layer. --A layer of wood or bark produced apparently during one growing p€triod; frequently, especially in woods of the temperate zones, divisible into early and late wood 

or bark (IAWA). 

Parenchyma.- -Tissue composed of cells that are typically brick -shaped or isodiametric and 

have simple pits; formed in wood from (a) fusiform cambial initials by later transverse 

divisions of the daughter cells (axial parenchyma), or (b) ray initials (ray or radial 

parenchyma). Syn. Soft tissue, Storage tissue. See also Parenchyma cell, fusiform. 

Note: Primarily concerned with the storage and distribution of food materials. Termed 

wood parenchyma or xylem parenchyma if occurring in the xylem, and phloem parenchyma if in the phloem. 

Parenchyma, apotracheal.- -Axial parenchyma typically independent of the pores or vessels. 

Note: This includes Terminal, Diffuse, and Banded apotracheal parenchyma. 

Parenchyma, axial. --Parenchyma cells derived from fusiform cambial initials. Syn. Longitudinal parenchyma (deprec.), Vertical parenchyma (deprec.), cf. Parenchyma, ray. 

Parenchyma, diffuse. --Single apotracheal parenchyma strands or cells distributed irregularly 

among fibres, as seen in cross section (IA WA modi£.). 

Parenchyma, paratracheal. --Axial parenchyma associated with the vessels or vascular tracheids (IAWA modi£.). Note: This includes Scanty paratracheal, Vasicentric, Aliform 

and Confluent parenchyma. 

Parenchyma, ray. --Parenchyma composing the rays wholly or in part. (IAWA modi£.). Syn. 

Radial parenchyma. 

Parenchyma, terminal. - -Apotracheal parenchyma cells occurring either singly or forming a 

more or less continuous layer of variable width at the close of a season's growth. 

Note: Before a distinction was made between "terminal" and "initial" parenchyma, 

this term was used to include both forms and is still used in this sense as a term of 

convenience. 

Perforation, simple. --A single and usually large and more or less rounded opening in the per-

£oration plate, cf. Perforation, multiple (IA WA). 

4 Excerpts from Committee on Nomenclature, International Association of Vvood Anatomists. International Glossary of Terms Used in Wood Anatomy. 1957. 

69 
Perforation, vessel.- -An opening from one vessel member to another (IAWA). 

Perforation plate. --A term of convenience for the area of the wall (originally imperforate) 

involved in the coalescence of two members of a vessel (IAWA). 

Perforation plate, scalariform. --A plate with multiple perforations elongated and parallel. 

The remnants of the plate between the openings are called Bars (IAWA). 

Pit. --A recess in the secondary wall of a cell, together with its external closing membrane; 

open internally to the lumen. Note: Essential components are the pit cavity and the 

pit membrane (IAWA modi£.). 

Pore. --A term of convenience for the cross section of a vessel or of a vascular tracheid 

(IAWA). 

Pore, solitary. --A pore completely surrounded by other elements (IAWA). 

Pore chain. --A series or line of adjacent solitary pores (IAWA). 

Pore multiple. --A group of two or more pores crowded together and flattened along the lines 

of contact so as to appear as subdivisions of a single pore (IA WA). Note: The most 

common type is a Radial pore multiple, in which the pores are in radial files with 

flattened tangential walls between them. Another type is a Pore cluster, in which the 

grouping is irregular. --

Protoplast.- -The mass of protoplasm enclosed by a cell wall. 

Ray.- -A ribbonlike aggregate of cells formed by the cambium and extending radially in the 

xylem and phloem (IA WA modi£.). Note: The terms Medullary ray and Pith ray are 

now restricted to the parenchyma connecting the primary cortex with the pith-.-

Ray, multiseriate.- -A ray two or more cells wide as seen in tangential section. 

Ray, uniseriate. --A ray one cell wide as seen in tangential section. 

Ray cell, procumbent.- -A ray cell with its longest axis radial (IAWA). 

Ray cell, upright. --A ray cell with its longest dimension axial (IAWA modi£.). Note: Such 

cells compose certain uniseriate rays and parts, typically the margins, of some 

multi seriate rays. 

Ring, annual.- -In wood and bark, a growth layer of 1 year as seen in cross section (IA WA 

modi£.), ct. Ring, growth. 

Ring, growth. --In wood and bark a growth layer as seen in cross section (IAWA modi£.). 

Spiral thickening. --Helical ridges on the inner face of, and part of, the secondary wall (IAWA). 

70 

Note: Often erroneously called tertiary spirals to distinguish them from the spirals of 

primary xylem. 
Storied (storeyed). --A term applied to the axial cells and rays in wood when these are arranged 

in horizontal series on tangential surfaces. Note: The term is applied to particular 

tissues, e.g., "storied parenchyma" or used in a general sense, as in "woods with 

storied structure." The presence of storied structure is the cause of the ripple marks 

visible with the unaided eye. 

Tannins. --A heterogeneous group of phenol derivatives; they appear as yellow, red, or brown 

substances within the cell, commonly in the vacuoles, but also within the cytoplasm.·:: 

Tracheid. --An imperforate wood cell with bordered pits to congeneric elements (IAWA modi£.)~ 

Vessel.- -An axial series of cells that have coalesced to form an articulated tubelike structure 

of indeterminate length; the pits to congeneric elements are bordered (IA WA modi£.). 

Syn. Trachea. 

Vessel member or element. --One of the cellular components of a vessel (IA WA). Syn. Vessel 

segment (deprec.). 

Wood.- -The principal strengthening and water -conducting tissue of stems and roots. Characterized by the presence of tracheary elements. Syn. Xylem. 

Wood, diffuse -porous.--Wood in which the pores are of fairly uniform or only gradually changing size and distribution throughout a growth ring (IA WA). 

Wood, early.- -The less dense, larger celled, first formed part of a growth ring (IAWA). Syn. 

Spring wood. 

Wood, late. --The denser, smaller celled, later fanned part of a growth ring (IAWA). Syn. 

Autumn wood (deprec.). cf. Summer wood. 

Wood, ring-porous. --Wood in which the pores of the early wood are distinctly larger than 

those of the late wood and form a well-defined zone or ring, cf. Wood, diffuse -poTous 

(BSI modi£.). 

Xylem. --See Wood. 

5 Description from Esau, Katherine. Plant anatomy, 2d ed. New York: John Wiley & 

Sons, Inc. 1 965. 

71 
REFERENCES 

1. Carlquist, S. 1958. "Wood anatomy of Heliantheae (Compositae)," Trop.Woods. 108: 1-30. 

2. Chattaway, M. Margaret. 1932. "Proposed standards for numerical values used in describing woods," Trop. Woods. 29: 20-29. 

3. Committee on Nomenclature, International Association of Wood Anatomists. 1957. "International glossary of terms used in wood anatomy," Trop. Woods. 107: 1-36. 

4. Doorenbos, J. 1965. "Juvenile and adult phases in woody plants." In: Encyclopedia ~ 

Plant Physiology XV I 1. New York: Springer-Verlag. 

5. Esau, Katherine. 1965. Plant Anatomy, 2d ed. New York: John Wiley & Sons, Inc. 

6. Hill, Ralph R. 1964. Key to identification of Rocky Mountain shrubs by characteristics of 

the wood aii:dbark. Denver: R. R. Hill, 3535 Benton Street. 9 pp. (mimeo). 

7. Hitchcock, A. S. 1925. Methods of Descriptive Systematic Botany. New York: John Wiley 

& Sons, Inc. 

8. Ingle, H. D., and Dadswell, H. E. 1956. "The anatomy of the timbers of the southwest 

Pacific area. IV. Cunoniaceae, Davidsoniaceae, and Eucryphiaceae," 

Austral.l. Bot. 4: 125-151, 

9 . Jane, Frank W. 19 56 . The Structure of Wood. London: A. & C . Black . 4 27 pp . , ill us . 

10. Larson, Philip R. 1964. "Some indirect effects of environment on wood formation," In: 

The Formation of Wood in Forest Trees. New York: Academic Press. 

11. Lawrence, George H. M. 1951. Taxonomy ~f Vascular Plants. New York: The MacMillan 

Co. 

12. Metcalfe, C. R. , and Chalk, L. 1950. Anatomy of the dicotyledons. Oxford: Clarendon 

Press. 

13. Panshin, Alexis John. 1964. Textbook 9! Wood Technology, 2d ed. New York: McGrawHill. 

14. Record, S. J. 1934. Identification of the timbers of temperate North America. New York: 

John Wiley & Sons, Inc.- - - --

15. Record, S. J., and Chattaway, M. Margaret. 

classifying dicotyledonous woods," 

1939. "List of anatomical features used in 

Trop. Woods. 57: 11-16. 

16. Reinders-Gouwentak, Cornelia. 1965. "Physiology of the Cambium and other secondary 

meristems." In: Encyclopedia of Plant Physiology XV I 1. New York: SpringerVerlag. 

17. Tiemann, Harry D. 1951. Wood Technology; constitution, properties and uses. 3d ed. 

New York: Pitman Pub. Corp. 

18. Whaley, W. G. 1965. "The interaction of genotype and environment in plant development." 

In: Encyclopedia of Plant Physiology XV I 1. New York: Springer-Verlag. 

72 
SPECIES INDEX 

Acer glabrum Torr.---------------------------------- 5, 36-37 

Alnus incana (L.) Moench----------------------------- 5, 42-43 

A. sinuata.{Regel) Rydb. ------------------------5, 42-43,46-47 

Amelanchier alnifolia Nutt. ---------------------------- 6, 12-13 

Arctostaphylos uva-ursi (L.) Spreng.------------------- 5, 48-49 

Artemisia tridentata Nutt. ----------------------------- 4, 8-9 

Betula glandulosa Michx. ------------------------------ 5, 44-45 

~· occidentalis Hook.--------------------------------- 5, 44-45 

Ceanothus sanguineus Pursh --------------------------- 6, 54-55 

C. velutinus Doug!. ---------------------------------- 6, 54-55 

Cercocarpus ledifolius Nutt. -------------------------- 6, 12-13 

Chrysothamnus nauseosus (Pall.) Britt. ----------------- 4, 10-11 

_Q. viscidiflorus (Hook.) Nutt. ------------------------- 4, 10-11 

Cornus stolonifera Michx. ---------------------------- 5, 46-47 

Crataegus douglasii Lind!. ---------------------------- 6, 14-15 

Holodiscus discolor (Pursh) Maxim. -------------------- 4, 14-15 

Juniperus communis L. ------------------------------- 4, 8-9 

]_. scopulorum Sarg. ---------------------------------- 4, 8-9 

Ledum glandulosum Nutt. ----------------------------- 5, 48-49 

LO!iiC"era involucrata (Rich.) Banks--------------------- 6, 58-59 

L. utahensis Wats. ----------------------------------- 6, 58-59 

Menziesia ferruginea Smith --------------------------- 5, 50-51 

Pachistima myrsinites (Pursh) Raf.--------------------- 6, 64-65 

Philadelphus lewisii Pursh ----------------------------- 4, 26-27 

Physocarpus rnaiYaeeus (Greene) Kuntze ---------------- 6, 16-17 

Populus tremuloides Michx. --------------------------- 5, 38-39 

P. trichocarpa T. and G.------------------------5, 38-39, 40-41 

Prunus emarginata (Doug!.) Walpers ------------------- 4, 16-17 

~giniana L. ------------------------------------- 4, 18-19 

Purshia tridentata (Pursh) DC. ------------------------- 6, 18-19 

Rhamnus alnifolia L'Her. ----------------------------- 6, 56-57 

~. purshiana DC. ------------------------------------ 6, 56-57 

Rhus g1abra L. -------------------------------------- 6, 52-53 

Ribesaureum Pursh ---------------------------------- 4, 28-29 

R. cereum Dougl. ----------------------------------- 4, 28-29 

R. hudsonianum Richards----------------------------- 4, 30-31 

~· irriguum Doug!.--------------------------- 5, 30-31,34-35 

~· lacustre (Pers.) Poir.----------------------- 4, 32-33, 34-35 

R. setosum Lindl.------------------------------------ 5, 32-33 !· viscosissimum Pursh------------------------------ 5, 34-35 

Rosa gymnocarpa Nutt. ------------------------------- 4, 20-21 

~· woodsii Lind!.------------------------------------ 4, 20-21 

Rubus idaeus L. ----------- ------------------------ -- 4, 22-23 

~· leu~mis Dougl. ------------------------------- 4, 22-23 

~· parviflorus Nutt.---------------------------------- 4, 24-25 

Salix scouleriana Barratt.----------------------------- 5, 40-41 

saffibucus cerulea Raf. ------------------------------- 5, 60-61 

~· racemosa L. -------------------------------5, 60-61, 62-63 

Shepherdia canadensis (L.) Nutt.----------------------- 6, 64-65 

Sorbus scopulina Greene------------------------------ 6, 24-25 

Spiraea betulifolia Pall.------------------------------- 6, 26-27 

Symphoricarpos albus (L.) Blake----------------------- 5, 62-63 

Taxus brevifolia Nutt. -------------------------------- 4, 8-9 

Vaccinium membranaceum Dougl. --------------------- 5, 50-51 

V. scoparium Leiberg-------------------------------- 5, 52-53 

73 

Dale , Arlene. 

1968. Comparative wood anatomy of some shrubs native 

to the Northern Rocky Mountains. U.S. Dep. Agr. , 

Forest Serv. , Intermountain Forest and Range Exp. 

Sta., Ogden, Utah, 84401. 73 pp., illus. (U.S. ForestServ. Res. Pap. INT-45) 

This paper describes some xylem characteristics of the 

more important shrub species of the Northern Rockies and 

presents a key for identifying shrub-wood specimens by 

microscopic characters 0 The paper contains photomicrographs of 55 shrub woods. 

Dale , Arlene . 

1968. Comparative wood anatomy of some shrubs native 

to the Northern Rocky Mountains 0 U.S. Dep. Agr 0 , 

Forest Serv. , Intermountain Forest and Range Exp. 

Sta., Ogden, Utah, 84401. 73 pp., illus. (U.S. Forest Serv. Res. Pap. INT-45) 

This paper describes some xylem characteristics of the 

more important shrub species of the Northern Rockies and 

presents a key for identifying shrub-wood specimens by 

microscopic characters. The paper contains photo micrographs of 55 shrub woods. 

Dale, Arlene. 

1968. Comparative wood anatomy of 

to the Northern Rocky Mountains 

Forest Serv. , Intermountain For

Sta., Ogden, Utah, 84401. 73 pp

est Serv. Res. Pap. INT-45) 

This paper describes some xylem ch

more important shrub species of the N

presents a key for identifying shrub-

microscopic characters. The paper c

graphs of 55 shrub woods. 

Dale , Arlene. 

1968. Comparative wood anatomy of 

to the Northern Rocky Mountains 

Forest Serv. , Intermountain For

Sta., Ogden, Utah, 84401. 73 pp

est Serv. Res. Pap. INT-45) 

This paper describes some xylem ch

more important shrub species of the N

presents a key for identifying shrub-

microscopic characters. The paper c

graphs of 55 shrub woods. 
Headquarters for the Intermountain Forest and Range Experiment Station are in Ogden, Utah. Project headquarters are 

also at: 

Boise, Idaho 

Bo;:eman, Montana (in cooperation with Montana State 

University) 

Logan, Utah (in cooperation with Utah State University) 

Missoula, Montana (in cooperation with University of 

Montana) 

Moscow, Idaho (in cooperation with the University of 

ld:.~lw) 

Provo, Utah (in cooperation with Brigham Young University) 
FOREST SERVICE CREED 

The Forest Service of the U. S. Department of Agriculture is 

dedicated to the principle of multiple use management of the 

Nation's forest resources for sustained yields of wood, water, 

forage, wildlife, and recreation. Through forestry research, 

cooperation with the Stdes and private forest owners, and 

managenwnt of the National Forests and National Grasslands, 

it strives - as directed by Congress - to provide increasingly 

greater service to a growing nation.