I

71-18,301
SMITH, Hanley Kerfoot, 19*t0THE BIOLOGY, WILDLIFE USE AND MANAGEMENT
OF SUMAC IN THE LOWER PENINSULA OF MICHIGAN.
Michigan State University, Ph.D., 1970
Agriculture, forestry & wildlife

U n iv e r s ity M icro film s, A XEROX C o m p a n y , A n n A rb or, M ic h ig a n

THE BIOLOGY, WILDLIFE USE AND MANAGEMENT
OF SUMAC IN THE LOWER PENINSULA OF MICHIGAN

By
Hanley Kerfoot Smith

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Fisheries and Wildlife

1970

ABSTRACT
THE BIOLOGY, WILDLIFE USE AND HANAGEHENT OF SUMAC
IN THE LOWER PENINSULA OF MICHIGAN

By
Hanley Kerfoot Smith

The purpose of this study was to evaluate critically three similar
plants, s t a g h o m sumac (Rhus typhina T o m e r ) , smooth sumac (Rhus gtabva
L.) , and a hybrid sumac (Rhus typh-ina>glabra) , in regard to their biology,
their importance to wildlife and their potential use in wildlife manage­
ment programs.

These species were chosen for study because they were

heavily utilized wildlife food plants and because their abundance in
Michigan was believed to be threatened by recent changes in that area’s
agricultural and forest land-use practices.
The principal study area comprised four square miles located in
the Manistee National Forest in northern Lower Michigan (about 17 miles
west of Cadillac, Michigan).

This area was characterized by well drained

podzol soils, second—growth northern hardwoods, cold winters and mild
summers, with fairly evenly distributed precipitation.

Field and nur­

sery experiments were also conducted in southern Lower Michigan, near
the Michigan State University campus at East Lansing.

This region,

about 120 miles SSE of the study area, was characterized by podzolic
soils, oak-hickory and beech-maple forest associations, and a more
moderate climate than that of the study area.
November 1966, and terminated in August 1969.

The study began in

Hanley Kerfoot Smith
Taxonomic studies revealed that the sumac species on the study area
may have been a hybrid of smooth and staghorn sumac.

It was demonstrated

that the suspected hybrid was intermediate in key taxonomic characters
to smooth and staghorn sumac.'

Additionally, it was demonstrated that

cross fertilization can occur between smooth and staghorn sumac.

The

specimens on the study area were therefore designated as Rhus typhina>

glabra .
It was found that the four square miles of the study area contained
thirty-three separate concentrations of sumac, covering an area of 28.7
acres.

Virtually all of the sumac occurred on abandoned farmlands which

had been planted to red pine (Pinna resinosa ) within the past ten years.
The most common woody associates were Rubus allegheniensisf Fvagaria

virginiana, Quercuo rubra, Rhus copallina, Amelanahier arborea and Populus
tramuloidec.

The most common herbaceous species in sumac areas were

Antennaria plant agini folia, Anaphalis margaritacea, Asolepias syriaoa,
Phyoalis hetcrophylla, Rumex aoetoaella and Solidago oanaaonsis.

The

cryptogamic layer was characterized by the mosses Poly triown juniperinwn
and Ceratodon purpureuc.
Considerable differences were found in the productivity, in terms
of numbers of fruits and stems, among the various groups of sumac on
the study area.

Data gathered in the summer of 1968 by randomly placed

quadrats indicated that the 28.7 acres of sumac produced about 2,300
pounds (oven-dry weight)
stem tips.

of fruit and 400 pounds (oven-dry weight) of

Only fruits greater than 3 inches in length and the first

2.5 inches of each stem were considered in these estimations.
Forty-nine percent of the plants on the study area were one year
old, and ninety-five percent were five years old or less; the oldest
being 20 years old.

Sumac was demonstrated to approach maximum

Hanley K. Smith
productivity, in terms of numbers of fruits and stems, at about seven
years, but there were insufficient data to reach a conclusion about the
age at which productivity begins to decline.

Some fruits were produced

on two-year-old plants but significant fruit production did not begin
until the fourth year.
An extensive literature survey revealed that smooth and staghorn
sumac are important wildlife food species throughout their ranges.
These plants appear to be of most significance in the diets of white­
tailed deer (OdocoZZeua virginfanus) , cottontail rabbits

(SyZviZagus

florida>iuc) , sharp-tailed grouse (Pediocetes phaoianellno) and ruffed
grouse (Botiasa wnheZZuc) .

The plant was shown to be heavily browsed by

white-tailed deer on the study area from November through March.

In

the winter of 1968-69 approximately half of the stems on the study area
were browsed, totaling about 200 pounds

(oven-dry weight) of sumac

stems or about seven pounds of stems browsed per acre of sumac.
average stem length browsed was 2.A inches.

All of the sumac fruit on

the study area were browsed in the winter of 1968-69.
about 2,300 pounds

The

This totaled

(oven-dry weight) of fruit, or about 82 pounds of

fruit of sumac browsed per acre.

When a sumac fruit was browsed,

the

entire fruit was eaten.
The stems and fruit of smooth, staghorn and

the hybrid sumac were

analyzed for nutrient composition, and the apparent digestibility

of

the

fruit of smooth sumac was determined.

Kuinac stems and fruits were

low

in crude protein, high in ether extract, and similar in gross energy

as compared on an oven-dry weight basis to three other Michigan deer
browses:

northern white cedar, jack pine, and big,-tooth aspen.

deer were fed a diet consisting only of smooth sumac fruits.

Six

The

digestibility data obtained indicated that sumac fruits were a good

Hanley K. Smith
energy source, but a poor source of protein, compared, on a dry-weight
basis, to sprays of northern white cedar.
It was found that sumac is rather easily grown from seed, and that
it may be transplanted with a high rate of success.

Mowing was also

demonstrated to be an acceptable method of reclaiming, as a deer browse,
clones which tiad exceeded the reach of deer.

ACKNOWLEDGEMENTS

It is a pleasure for me to extend my thanks to:
Dr. Leslie W. Gysel, Chairman of my Guidance Committee, for his
advice, encouragement and support throughout the study.
Drs. Gerhardt Schneider, Duane E. Ullrey and Donald P. White,
members of my Guidance Committee, for their assistance and
advice.
Dr. John H. Beaman, for his aid in the Identification of many
plant species.
Mrs. Betty L. Shoepke, for her technical assistance in performing
the nutritional analyses.
the personnel of the United States Forest Service, especially
Ronald Scott and George Irvine for their cooperation during
my field studies in the Manistee National Forest.
the personnel of the Houghton Lake Wildlife Research Station,
Michigan Department of Conservation, for their aid and
cooperation during the deer nutrition studies.
my wife, Ellie, for her constant support and encouragement during
the study and for aid in preparation of the manuscript.

ii

TABLE OF CONTENTS
Page
INTRODUCTION

................................

1

Study A r e a .................................................... 1
BIOLOGY OF THE P L A N T .................................................. 5
R a n g e ........................................................... 5
Site Requirements ...........................................

5

Taxonomy

....................................................

5

Phenology ....................................................

9

Reproduction, Root System andGrowth Form ..................

9

Diseases and Insect Infestations

..........................

Plant Associates, Productivity andAge Structure
SUMAC

..........

11
15

AS A WILDLIFE FOOD I T E M ...................................... 24
Survey of the L i t e r a t u r e .................................... 24
Availability

................................................ 27

U s e ............................................................ 28
S u m m a r y ........................................................32
THE PROXIMATE ANALYSIS AND APPARENT DIGESTIBILITY OF SUMAC

. . . .

Methods and M a t e r i a l s .................

35
35

R e s u l t s ........................................................39
D i s c u s s i o n ................................................... 44
S u m m a r y ........................................................48
ESTABLISHMENT AND PROPAGATION OF THEP L A N T .......................... 50
Collection and Germination of Seeds .........................

50

Propagation from S e e d ........................................ 52
iii

Page
Transplants.....................................................58
Propagation by Physical Disturbance.

. , .................

60

Summary..........

63

C O N C L U S I O N S ...................

65

REFER EN CES ................................................................ 66

APPENDIX.............................................

73

VITA.................................................................... 76

iv

LIST OF TABLES
Table
1

Page
Size and grouping of sumac concentrations on the
study area
............

15

2

Woody plants growing in association with sumac, as
determined by 180 randomly chosen milliacre plots........... 16

3

Frequency of herbs growing in association with sumac,
as determined by 100 randomly chosen milliacre plots . . .

17

4

Productivity of sumac groups on the study a r e a ..............19

5

Accuracy of estimating sumac age by counting the branch­
ing angles of plants 1 to 15 years o l d ......................21

6

Age profile of sumac plants on the study area, as
determined by 320 randomly chosen milliacre plots........... 22

7

The relationship of age to stem and fruit productivity
in s u m a c ......................................................23

8

Partial list of authors reporting use of sumac by
game s p e c i e s ................................................. 25

9

Frequency and density of woody plants emerging from a
14-inch snow cover In forest openings, as determined by
50 randomly selected 60 square foot plots (6* x 10')
sampled on the study area in the winter of 1967— 68 . . . .

27

10

Location of sumac clones used in analyses.................... 36

11

Proximate analyses of sumac stems and fruit, expressed
on a fresh weight basis.......................................40

12

Sumac browse and water intake, apparent digestibility
and apparently digestible energy during the last seven
days of Phase III............................................. 42

13

Urine and methane output, nitrogen balance, and appar­
ently metabolizable energy during the last seven days
of Phase I I I ................................................. 43

14

Weight losses during Phases I-III of the digestibility
study.................................

v

44

Table

Page

15

A comparison of the proximate analyses of several winter
browses.
(All analyses except dry matter are expressed
on an oven-dry weight basis.)................................ 45

16

A comparison of cedar sprays and sumac fruits in terms
of intake, apparent digestibility, apparently digestible
energy, and weight loss...................................... 47

17

Weight of sumac seeds per fruit and number of seeds
per pound..................................................... 52

18

The success of several seed planting trials at the
Tree Research C e n t e r .........................................56

19

Comparison of several transplant methods attempted at
the Tree Research C e n t e r ...........................

59

Productivity of sumac before and after mowing.

62

20

vi

. . . . . .

LIST OF FIGURES
Figure

Page

1

Map of the study area.....................................

2

2

Average monthly precipitation and temperature at
Cadillac, Michigan .......................................

4

A hybrid index, demonstrating the expression of the
taxonomic characters of Rhuo glabra , Rhuc typhina ,
and a suspected hybrid sumac ............................

8

3

4

5

Stem and fruit of Rhus glabra , Rhus typhina, and their
hybrid Rhus typhina?glabra .......................

10

Diagrammatic drawings of the root systems of five
sumac c l o n e s ...................

12

6

Stem and foliage of a normal sumac plant and a sumac
plant infested with the mite Erlophyes r h o t s ............. 14

7

The monthly consumption by deer of sumac stems on the
browse plots from May 1967 through April 1969...........

30

Rate of disappearance of sumac fruit susceptible to
deer browsing, on the study area, in the autumn and
winter of 1967-68 and 1968-69............................

33

Percentage of sumac seeds germinating after various
time intervals in concentrated sulfuric acid . . . . . .

53

Rate of germination of seeds scarified in sulfuric
a c i d .....................................................

54

Percentage of seeds germinating when planted at various
depths in soil trays, under greenhouse conditions. . . .

57

8

9

10

11

vii

INTRODUCTION

The purpose of this study was to evaluate critically three similar
plants, staghorn sumac (Rhus typhi-na Torner), smooth sumac (Rhus glabra
L.), and a hybrid sumac (Rhus typ1vlna>glabra) , in regard to their biology,
their importance to wildlife and their potential use in wildlife manage­
ment programs.

These species were chosen for study because they were

heavily utilized wildlife food plants and because their abundance in
Michigan was believed to be threatened by recent changes in that area's
agricultural and forest land-use practices.
The study is presented in four sections:

the general biology of

the plant, its use by wildlife, its nutritional value, and nursery and
establishment techniques.

Although the topics are discussed separately,

each is meant to complement the others.

Hopefully, this study will pre­

sent a perspective which wildlife biologists and foresters may use in
evaluating the importance of sumac in their land management programs.

Study Area
The principal study area comprised four square miles

(T21N-R12W,

sections 3, 4, 9 and 10) in the Manistee National Forest in Wexford
County in northwestern lower Michigan (Figure 1).

Characteristic topo­

graphic features of this region are hilly moraines dissected by small
streams.

The sumac on the study area is found on Blue Lake and

Kalkaska sands.

Both soils are well drained podzols developed on glacial

drift, originally supporting northern hardwoods, hemlock and white pine.

Figure 1.

Map of the study area.

2

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3
Blue Lake soils differ from Kalkaska soils in that the former possess
a Bt horizon and fine textural bands in the subsoil (personal communi­
cation, H. L. Weber, U. S. Soil Conservation Service, Traverse City,
Michigan).
The climate of this region of Michigan is characterized by cold
winters and mild summers, with fairly evenly distributed precipitation.
Climatological data from the U. S. Weather Bureau at Cadillac,
east of the study area, are summarized in Figure 2.

18 miles

The area averages

110 freeze-free days per year, with the last freeze usually occurring
between May 20 and May 30 and the first freeze usually occurring between
September 10 and September 20 (Eichmeier et at, , no date).

Average

annual precipitation is 30.8 inches; 8.4 Inches being the water equiva­
lent of the 60.2 inches of annual snowfall.
Of primary interest in this study were the abandoned farmlands on
which the sumac was found.

These openings comprised about seven percent

of the area, the remainder being medium to well stocked stands of aspen
and northern hardwoods.
trees in the area:

Berner (1969) listed the following as common

American elm, black ash, white ash, red maple, sugar

maple, big-toothed aspen, quaking aspen, basswood, red oak and black
cherry.

Ubiquitous in the forest openings were plantations of red

pine, white pine, and white spruce.

These plantations, which covered

nearly 400 acres, had been established since 1961.
Many field and nursery experiments with sumac were conducted in
Ingham, Shiawassee and Clinton Counties near the Michigan State Uni­
versity campus at East Lansing.

This region, about 120 miles SSE of

the study area, is characterized by a more moderate climate than that
of the study area, podzolic soils, and oak-hickory and beech-maple
forest associations.

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60

TEMPERATURE

50-

AO

30

20

JA N

FEB

MAR

A PR

MAY

JU N

JU L

MONTHS

Figure 2. Average monthly precipitation and temperature at
Cadillac, Michigan (Climatological Data, Michigan, 1928-1962).

BIOLOGY OF THE PLANT

Ran Re

Smooth sumac is native throughout southern Canada, and all of the
48 contiguous states, except California.

Staghorn sumac occurs naturally

from Nova Scotia south to North Carolina, and west to Minnesota and
Iowa (Barkley, 1937; West and Arnold, 1956).

Additionally, these

plants have been widely introduced in the United States and southern
Canada as ornamentals and for erosion control (Boyd, 1943b).

Staghorn

sumac is cultivated in Europe and Asia where it is a source of tannin
(Baczuk and Bukiewicz, 1961; Quraishi et at., 1964).

Site Requirements

Both species are pioneer shrubs or small trees usually found in
open areas on well drained sands and sandy loams.

Typical sites include

abandoned fields, roadsides, railroad rights-of-way, fence rows, burned
or denuded areas, and young forest plantations (Bingham, 1937; Boyd,
1943b; Clements, 1920; Dice, 1923; Hirth, 1959; Verts, 1957).

Smooth

sumac is characteristic of the forest-prairie ecotone of the midwestern
United States (Bray, 1960; Ewing, 1924; Shelford and Winterringer,
1959) , and both species are common on the abandoned farmlands of the
northeastern United States and southeastern Canada (Hirth, 1959).

Taxonomy

Rhus typh-Lna and Rhus gtabva are quite similar morphologically,
the principal differences being that the fruit and stem of the former
5

6
are densely pubescent, while those of the latter are glabrate or nearly
so.

Several authors have suggested that they may hybridize, but this

has never been proved (Barkley, 1937; Green, 1906; Little, 1945;
Sargent, 1891).

Both species are polymorphic and have been subjected

to considerable nomenclatural subdivision.

As a result, three names,

which may represent hybrids of Rhus typhi-na and Rhus glabra, remain in
the literature today:

Rhus pulvinata Green, Rhus boreali-s Green, and

Rhus glabra var. borealis.
hybrid

and was described

The sumac species on the study area was a
adequately by any of these latter names.

Two approaches were used to determine the taxonomic position of
the sumac plants on the study area.

The first was cross-fertilization

between staghorn and smooth sumac and the second was the development
of a hybrid index.
In June of 1969, ten pistillate flowers of Rhus typhina* and ten
pistillate flowers of Rhus glabra* were covered with white, waterresistant paper bags prior to their blooming.

When the flowers of each

began to open, they were dusted with the pollen of the opposite species.
The lower flowers of the inflorescence opened first, with about a sixday period between the first and last bloom on each stem.

The flowers

were dusted each day that they were observed to be in bloom, and the
stems were unbagged only during the few moments that were required to
dust them.

These plants are believed to be insect pollinated (Heimsch,

1940) and care was taken that they not be contaminated by insects
during the dusting process.

Viable seed stock has been obtained from

the crosses, Indicating that the two species can hybridize.

*The experimental clones were located in Ingham County,
T3N-R1W-S6, and vouchers of each clone were deposited in the Beal
Herbarium, Michigan State University,

7
The hybrid index, as discussed by Anderson (1949) and Rollins
(1957), provides a semiquantitative method of ranking suspected hybrids.
Contrasting characters are chosen on the parent species and these
characters are given numerical scores, with an intermediate score pro­
vided for characters that are common to both.
the scores of several specimens are summed.
were chosen for smooth and staghorn sumac:

An index results when
Five contrasting characters

(1) the length of the

pubescence on the fruit,

(2) the presence or absence of an abscission

zone on the fruit stalk,

(3) the density of the pubescence on the

stem,

(4) the length of the pubescence on the stem, and (5) the density

of the pubescence on the petiole.

Seventeen specimens identified as

Rhus typhina and ten identified as Rhus glabra from the Beal Herbarium
at Michigan State University were scored and used as standards for
their species.

Forty specimens, randomly collected on the study area,

were also scored.

The results appear in Figure 3.

According to the

descriptions of Barkley (1937), the monographer of the genus Rhus3

Rhus glabra would receive a score of seven or less and Rhus typhina
would have a score of 16 or more.

The names assigned to some of the

museum specimens, those with intermediate scores, did not correspond
to their accepted descriptions, indicating that they were hybrids that
had been misidentifled.

The characters of the specimens collected on

the study area appear to be intermediate to those of Rhus typhina and

Rhus glabra.

The forty specimens from the study area had a mean score

on the hybrid index of 13.4 and were thus judged to demonstrate a ten­
dency toward greater expression of the characters of Rhus typhina.
These observations suggest that the plants on the study area are
a hybrid swarm.

The symbol ”>" has been suggested by Li (1957) to

10“

6-

NUMBER

OF

SAM PLES

o

4-

□

O

□
2

-

A
I
10

I

n

0
□

O

r
12

I
13

14

15

16

T
17

n
18

o
o
i

i

i

19

20

21

r
22

CUMULATIVE SCORE
Rhus typhina O
Rhus glabra A
Rhus hybrid □

ItfDEX CRITERIA
A.

Length of hair on
fruit (ram.)
Score
1
< 0.25
2
0.25 - 0.50
0.50 - 1.00
3
1.00 - 1.50
4
1.50 - 2.00
5
2.00+
6

Abscission zone on
fruit stem
Score
1
obvious
obscure
2
none
3

C.

Density of hair on
stem (mra.)
Score
1
none
2
scant
medium
3
heavy
4

D.

Length of hair on
Score
stem (mm.)
no hair
1
2
< 0.50
0.50 - I.00
3
4
1.00 - 1.50
1.50 - 2.00
5
6
2.00+

E.

Density of hair on
Score
petiole
1
none
2
scant
medium
3
4
heavy

Figure 3. A hybrid index, demonstrating the expression of the taxonomic characters
of Rhus glabra, Rhus typhina, and a suspected hybrid sumac.

00

9
indicate the probable direction of gene flow in an introgressant popu­
lation.

From the information available, I have designated the sumac

population on the study area as Rhus>typhina glabra (Figure 4).

Phenology
Sumac in southern lower Michigan remains dormant until about the
first week in May, when bud elongation occurs.

Rhus glabra and Rhus

typhina flower concurrently during the latter part of June and the first
two weeks of July.
by mid-September.

The first fruits appear in mid-July and are ripe
The phenological events of the sumac on the study

area occur about one week later than the corresponding events in Ingham
County, 120 miles to the south.

Gilbert (1961) has fully documented

the phenology of sumac in southern Michigan.

Reproduction, Root System and Growth Form

Although sumac plants produce an abundance of seeds, reproduction
occurs primarily by root sprouts.

The root system spreads laterally

from the center, frequently branching and sending up shoots.

Duncan

(1935) found that Rhus copallina roots spread outward at a rate of
about 24 inches per year.

Gilbert (1959) reported that the annual

spread of several smooth and staghorn sumac clones varied from 27 to
76 inches.

Sumac roots are primarily shallow, but some plants have

been found to send tap roots as deep as 90 inches (Weaver, 1919).
Fifteen small clones were excavated on the study area in August
of 1967.

Most of the roots were found from just below the surface

litter to a depth of about 4 inches, but a few were traced to depths
of almost 36 inches.

In all cases some part of the root had been

severed by a plow when the area was planted to pine.

Five clones

10

Rhus typhina

Rhus typhina>glabra

Rhus glabra

Figure 4. Stem and fruit of Rhus typhina , Rhus glabray and their
hybrid Rhus typhina >glabra (X 2/3).

11
which are considered illustrative are diagrammed in Figure 5.
arise from swollen nodes along the roots.

Stems

The fact that new stems may

arise from dormant root system is demonstrated by the presence of
long-dead aerial portions on the same root as one—year— old plants.
Several of the older, rotten nodes had been colonized by ants.
The aerial portions of an undisturbed sumac clone are character­
istically pyramidal with the oldest and tallest stems in the center.
The stems sprout centrifugally until the clone is about 15 years old,
at which time it loses vigor and the spread of new stems slows, while
the older center stems die (Gilbert, 1966).

In contrast, the sumac

stems on the study area do not demonstrate a consistent size structure
because virtually all of the clonal root systems have been fragmented
by a pine planting plow.

The possibilities of propagation by root

cutting are discussed briefly in a later section.

Diseases and Insect Infestations
Sumac is of limited economic importance in the United States, and
consequently reports of its diseases are rather uncommon in the litera­
ture .

Judd (1963) reported infestations of the red pouch gall

Metophis phots on sumac in Ontario.

Pirone et at.

(1960) noted that

several species of sumac are known to be infected by fungi of the
genera Ptteotapta, Fusaptum^ Cpyptodtnpopthe, Physatospora, Veptt-

atZZtim^ and Sphaepotheca. Heavy infestations of the chalcld fly
Idtomaopomepus btmtauZupemis were found in smooth sumac fruits by
Lovell (1964) in Kansas.

Lovell also noted that a fungus of the genus

Pythtwn commonly infected sumac seedlings raised in the laboratory.
In the summer of 1968, nearly 20 percent of all one-year-old
stems on the study area were infested with the mite Eptophyes phots

12

Irr

cut

£_!£.

'Jfr

*ir

1v 1__ J£i

lyr

io“

cut

2yvi

KIT
Llv* root*
Llv* i t * s .
M 4d

Figure 5.
sumac clones.

it« * .

Diagrammatic drawings of the root systems of five

13
(Eriophyidae: Acarina).
tion of leaves,

The disease was characterized by the deforma­

the fleshy proliferation of tissues and the production

of extremely aberrant foliage (Figure 6).

The aerial portion of a

plant infested with this mite dies, and is not browsed by deer.

In

some parts of the study area this disease approached a 100 percent
infestation.

I have also noticed this disease on museum specimens of

Rhus typhina from New Hampshire and Massachusetts.
The larval stage of the moth Hotcocera chaZcofronteZZa (Blastobasidae: Lepidoptera) was observed to cause extensive damage to the
fruit of staghorn sumac.

Eight of ten randomly selected clones in

Ingham County had sustained damage to more than 90 percent of their
fruits from this insect in 1969.

The larvae were found to be active

in the fruits during the months of September and October, eating the
fruit coats of the inner fruits, and leaving the inflorescences appear­
ing outwardly undamaged.

This insect and the mite previously discussed

were identified by Dr. W. E. Wallner of the Entomology Department at
Michigan State University.
It is believed that this is the first report of members of the
genus Rhus being infested by either HoZcocera chaZcofronteZZa or

Eriophyes rhoi-s.

However, E. rhois has been reported to infest another

species of the family Anacardlaceae, Toxicodendron radZcans (Felt,
1940) .
Considerable problems resulted from the girdling of the hypocotyl
of seedlings which were planted in unsterile soils in the laboratory.
The disease was believed to be caused by a "damping off" fungus of the
genus PythZum , but this was not verified.

The girdling was easily pre­

vented by dusting the seeds and seed bed with Captan, a commercial
fungicide.

Figure 6.
Stem and foliage of a normal sumac plant (r) and a
sumac plant infested with the mite Eriophyes rhois (1).

15
Plant Associates, Productivity and Age Structure
In the summers of 1967 and 1968, the four square miles of the
study area were thoroughly searched in order to locate the main sumac
areas.

Thirty-three separate sumac concentrations were found on the

study area and these are indicated on Figure 1.

For convenience,

these separate concentrations are combined into five groups

(A through

E) on the basis of their proximity and the similarity of sites

Table 1.

Group

Size and grouping of sumac concentrations on the study area

Area
Number

Size
(ft2)

1
2
3
4
5
6
7
8
9
10
11

2,300
9,000
10,400
3,800
5,800
2,400
13,500
3,700
16,400
6,500
6,500
80,200

A

Total

Group

Area
Number

Size
(ft2)

C

14

409,800
409,800

15
16
17
18
19
20
21
22
23

5,600
25,000
12,600
68,900
7,500
10,200
4,800
26,000
6,800
167,400

Total

D

Total

B

12
13

Total

(Table 1)

Group

Area
Number

Size
(ft2)

24
25
26
27
28
29
30
31
32
33

12,500
9,200
5,000
5,400
6,300
11,200
7,400
6,200
5,500
7,800
76,500

E

Total

Total sumac area
in acres = 28.72

3,100
514,500
517,600

Groups A, D and E were each sampled by 60 milliacre plots, while
Groups B and C were each sampled by 70 milliacre plots.
square and randomly placed within each sumac area.

The plots were

The frequency and

number of all woody species were recorded for each plot.

The frequency

by species of all herbaceous plants was noted in 100 plots randomly

16
chosen from the total run.

All sumac plants in the plots were aged

and their height and stem lengths measured.

Additionally, the numbers

of fruiting, non-fruiting and diseased stems were tallied.

Only those

plants rooted in the plot w e r e .considered in any of the tallies.
The cryptogram layer in the sumac areas was measured by 19 randomly
placed 60-foot line intercepts.

The 60-foot line was composed of two

perpendicular 30-foot lines crossed at the midpoint of each line.

Only

that portion of the ground cover actually traversed by the line was tallied.

Plant Associates:

The frequency and density of woody plants grow­

ing in the sumac areas are presented in Table 2.

Table 2.

Rubus altegheniensis

Woody plants growing in association with sumac, as determined
by 180 randomly chosen milliacre plots"**

Stems/
Acre

Frequency
(Percent)

Species

Rubus aiiegheniensis
Firms reoinosa
Querous rubra
Rhus aopa IU n a
Amelanahier arborea
Populus tremuloides
Prunus serotina
Ulmus amerioana
Prunus virginiana
Fraxinuo amerioana

Percent of plots with no woody plants except sumac:
+ Source of botanical nomenclature:
*Not counted

*
261
44
311
22
27
11
16
5
5

22.2
26.1
2.7
2.2
2.2
1.6
1.1
0.5
0.5
0.5
49.4

Gleason and Conquist, 1963

was by far the most common woody species, often exceeding 50 plants per
plot, but because of time considerations the actual number of Rubue was
not determined.

Firms resinosa, representing stock that had been

planted during the previous ten years, was the next most common woody

17
species.

The eight remaining shrubs and trees were rather infrequent

on the sumac areas and rarely reached a height of three feet there.
Almost half of the plots were completely devoid of any woody species
except sumac.

The herbaceous layer consisted of 24 species, the most

conspicuous of which were Asclepias syriaca and Solidago canadensis
(Table 3).

Table 3.

The cryptogamic layer was characterized by the mosses

Frequency of herbs growing in association with sumac, as
determined by 100 randomly chosen milliacre plots'*-

Species

Antennaria plantaginifolia
Anaphalis margaritaaea
Asclepias syriaca
Physatis heterophylla
Fragaria virginiana
Rwnex acetoaella
Solidago canadensis
Hypevnicwn punctatum
Ambrosia artemisiifolia
Erigeron canadensis
Hieracium aurantiacum
Prunella vulgaris
Tragopogon dvtbius
Solidago sp.
Ranunculus septentrionalis
Anemone aylindrica
Pteridium aquilinum
Hieracium gronovii
Hieracium florentium
Apocynum cannabiutn
Erigeron annuus
Centaurea maculosa
Achillea millefolixw\
V i d a cracca

^Source
of botanical nomenclature:
JL
Treated with herbaceous layer

Frequency
(Percent)

74
70
55
35
28
27
27
21
18
14
12
12
11
7
6
5
3
2
2
1
1
1
1
1

Gleason and Cronquist, 1963

18

Polytriaum jun'Lperinum and Ceratodon pwrpuveus.

The most common lichen

was cladonia arbuscula » followed in descending order by C. ahlorophaea*

C, pyxidata and c. oristatella •

The ground cover was found to average

four percent bare soil, fourteen percent mosses and lichens, and eightyone percent grasses, litter and herbs.

Productivity:

A compilation of the sumac productivity data col­

lected in August of 1968 is presented in Table 4.

Group A was the least

productive area, perhaps because it was the only area that had not been
disturbed by the pine planters’ plow.
largest and most homogeneous areas.

Groups B and C were by far the
Group B was also the most product­

ive group, possibly because it was the only area with a predominantly
southern aspect.

It should be noted, however,

that the effect of slope

on sumac growth was not analyzed in this study.
A high proportion of the stems in Groups A, B and C were infested
with the mite Eviophyes rhois , while relatively few of the stems in
Groups D and E were diseased.

Ninety-eight percent of the infested

stems were one year old, and the remaining two percent were two years
old.

No explanation was found for the discrepancies in disease occur­

rence by group or age.
Considerable differences were seen between the per acre weights of
stems and fruits of the various groups.

These differences probably

reflect the density, vigor and age structure of the sumac populations
represented.

Additionally,

the differences seen between fruit weights

could be a result of an unequal distribution of pistillate clones.

In terms of wildlife management, fruit and stem production are most
important.

The data in Table 4 indicate that the 28.7 acres of sumac on

the study area in 1968 produced about 2,300 pounds

(oven dry weight) of fruit

Table 4.

Group

Productivity of sumac groups on the study area in August of 1968

Acres

Number
of
Plots

%
Area
Sampled

Plants/1
Acre

Browse2
Stems/
Acre

Diseased3
Stems/
Acre

Good4
Fruits/
Acre

Poor5
Fruits/
Acre

Total5
Stems/
Acre

Weight/7 Acre
(lbs.)
a) Fruit b) Stems

A

1.84

60

3.15

8,565

7,785

1,942

1,776

166

11,670

50

11

B

11.88

70

0.58

13,213

11,611

3,203

3,632

57

18,504

102

16

C

9.40

70

0.74

10,467

7,851

2,574

2,102

320

12,847

59

11

D

3.84

60

1.56

7,480

9,379

183

2,616

750

12,928

73

13

E

1.76

60

3.42

10,873

13,214

697

2,839

282

17,032

80

18

Plants - discrete aerial portions
2Browse stems - non-fruiting, healthy new growth
^Diseased stems - stems infested with mites
4Good fruits - normal fruits exceeding 3 inches in length
5Poor fruits - damaged or diseased fruits, or fruits less than 3 inches in length
6Total stems - the sum of all fruiting and non-fruiting stems
7Weight (oven dry) of: a) all good fruits, b) first 2.5 inches of all browse stems

20
and 400 pounds (oven dry weight) of stem tips.

Only fruits greater than

3 inches in length and the first 2.5 inches of each stem were considered
in these estimations.

Dalke and Spencer (1944) and Krefting et at .

(1955) have reported that deer may browse sumac plants so intensively
that the plant may be killed.

However, there were no indications that

browsing hindered productivity on the study area, as there appeared to be
no reduction in stem and fruit production on plants browsed the previous
year.

Age structure:

Sumac stems are straight and stout and new growth

arises from lateral buds, thereby forming an angle with the axis of the
old stem.

Employing this information, Gilbert (1959) found that the

plant may be aged by counting the greatest number of continuous branching
angles.

To determine the accuracy of this method, I estimated the age

of 264 plants by counting the branching angles and compared this estima­
tion with the age as determined by counting the annual growth rings of
those plants.

The annual ring count was assumed to represent the actual

age of the plant.

The results of these comparisons appear in Table 5.

No errors were made aging plants three years old and younger.

With plants

older than three years, the estimation of age was correct to within one
year in a high percentage of cases.

Since the great majority of the

plants fall into easily and accurately aged categories, the angle counting
method was used in all of the age determinations in this study.
The age profile (Table 6 ) indicated a young and vigorously growing
population which appeared to have begun expansion within the last ten

*0 ne fruit = 12.8 grams, oven-dry weight, based on a sample of
200 randomly chosen fruits.
One stem = 0.64 grams, oven-dry weight,
based on a sample of the first 2.5 inches of 200 randomly chosen stems.

21
Table 5.

Age
(years)

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15

Accuracy of estimating sumac age by counting the branching
angles of plants 1 to 15 years old

Sample
Size

Number
Correct

25
25
25
23
20
18
17
19
16
16
13
9
8
4
4

25
25
25
20
13
13
12
15
9
11
5
7
7
1
2

Departure of esti­
mate from actual age
H~1
+2
±3
(years)

3
6
4
4
1
5
4
5
2
1
3
2

1
1
1
2
1
2

1
2
1

+0

100
100
100
86
65
72
71
78
56
68
38
78
88
25
50

Percent
Correct
+1
+2
(years)

100
95
94
94
84
74
93
76
100
100
100
100

years from clones that had lost vigor or become dormant.

±3

100
100
100
95
74
100
92

100
100
100

In Groups B,

C and E, the beginning of the increase in growth rate corresponded
roughly to the dates on which pine was established in the area.

The

origins of Groups A and D were more obscure, but may have been related
to the movement of heavy machinery in 1958 and 1959 when Wagon Wheel
Road, which paralleled Group D, was repaired, and when a pine enclosure
was constructed near Group A.

Group D had an elevated percentage of

two-year-old plants, probably reflecting a response to the planting of
pine in this area in 1967.

The number of three-year-old plants appeared

to be lower than expected from the general trend of the age profile.
This inconsistency is probably best explained by the severe drought
that occurred during the growing season of 1966, thus lowering the
number of three-year-old plants.

In that year only 1,99 inches of rain

22
Table 6 .

Age profile of sumac plants on the study area, as determined
by 320 randomly chosen milliacre plots

Age Structure by Group
B
C
D
(Percent)

Age
(years)

A

1
2
3
4
5
6
7
8
9
10
11
12
16
20

64
14
10
9
2
1
P
P
X
x+
X
X
X
X

47
17
7
8
6
3
3
p*
P
P
P
X
X
X

61
13
6
8
5
2
1
1*
1
P
X
X
X
P

20
40*
13
16
5
2
1
1
P
P
X+
P
P
X

E

33
28
11
18
7
3*
P
P
X
X
P
X
X
X

Composite
(Percent)

49
21
9++
11
5
2
1
1
P
P
P
P
P
P

*Year in which pine was planted in area
+Year in which heavy equipment operated on area
++May be a result of severe drought in summer of 1966
P-Present in amounts less than one percent
X-Not present

fell at Cadillac

during the months of May and June, as compared with

average for this

period

the

of 8.92 inches.

Eleven of the sample plots were 100 percent shaded by overtopping
trees.
follows:

The age distribution of the 49 plants in these plots was as
one year old, 14%; two years old, 27%;

three years old, 21%;

four years old, 12 %; five years old, 18%; six years old, 4%; and seven
years old, 4%.

The trend here, as compared to the composited age struc­

ture of the area (Table6 ) was a failure to reproduce under shaded con­
ditions.

Such a lack of reproduction was often noted when sumac was

found growing under the semi-closed canopy of invading trees, and may
be the result of competition for light or water or both.

23
The relationship between age and the production of stems and fruits
is seen in Table 7.

The older plants approach maximum productivity at

about seven years, but there are insufficient data to reach a conclusion

Tab le 7.

The relationship of age to stem and fruit productivity in sumac

Age

Sample
Size
(Plants)

Browse
S terns

Browse
Stems/
Plant

Total
Fruit

Fruit/
Plant

1
2
3
4
5
6
7
8
9
10

1,620
688
292
375
171
66
33
26
15
5

1,615
815
372
496
290
96
77
63
29
10

1.00
1.18
1.27
1.32
1.69
1.45
2.33
2.42
1.93
2.00

5
96
102
306
237
93
45
22
24
5

.00
.14
.35
.82
1.39
1.41
1.36
.85
1.60
1.00

All
S terns*

1,620
911
474
802
527
189
122
85
53
15

Total
Stems/
Plant*

1.00
1.32
1.62
2.14
3.08
2.86
3.69
3.27
3.50
3.00

^Includes stems and fruit

about the age at which the plants begin to decline in productivity.
fruits are produced in the second year* contrary to the statements by
Boyd (1943b) and Spinner and Ostrom (1945) that sumac does not fruit
until the third or fourth year.

Some

SUMAC AS A WILDLIFE FOOD ITEM

Survey of the Literature

A partial list of references to the utilization of sumac as a
food item by wildlife appears in Table 8 .

This survey treats only game

species, and thus excludes the many songbirds and rodents that are known
to eat this plant (Martin et at ., 1951) .
Sumac fruits are eaten by many gallinaceous birds, but are believed
to be of major importance only in the diets of the ruffed grouse (Bump

et at. , 1947) and the sharp-tailed grouse (Ammann, 1957).

Although

several authors list sumac as a common food of the bob-white quail, it
has been demonstrated that it is of low energy value to that bird
(Errington, 1936; Newlon et at. , 1964), prompting researchers to sug­
gest that this fruit may be sought for a specific nutrient (Nestler
and Bailey, 1944),

The fruit is also commonly mentioned in lists of

wild turkey foods, but it seldom exceeds one percent of that animal's
annual diet (Korschgen, 1967).
Reports of heavy rabbit and squirrel use of sumac bark are usually
associated with deep snow conditions, suggesting that it is primarily
an emergency food for these species (Brown, 1947; Packard, 1956).

How­

ever, Hickie (1940) lists sumac among the preferred winter foods for
cottontail rabbits in Michigan.
Various sumac species are utilized by deer throughout much of the
United States.

Preference of sumac appears to be somewhat regional as

it is of secondary Importance to deer In the southeastern states
24

Table 8.

Partial list of authors reporting use of sumac by game species

Author (s)

Game
Species

Allen, R. H., Jr., and A. M. Pearson (1945)
Ammann, G. A. (1957)
Banasiak, C. F. (1961)
Brown, H. L. (1947)
Bump, G. et al. (1947)
Crispens, C. G. et al. (1960)
Dahlberg, B. L., and R. C. Guettlnger (1956)
Dalke, P. D. et al. (1946)
Errington, P. L. (1936)
Errington, P. L., and F. N. Hamerstrom, Jr. (1936)
Forbush, E. H. (1916)
Goodrum, P. D., and V. H. Reid (1962)

Cv
Tc,PP
Ov
Sf
Bu
Lc
Ov
Mg
Cv
Cv
Cv,Bu
Ov

Rg
Rg
Rsp
Rsp
Rsp
Rt,Rg
Rg

F
F
F-S
B-S
F
F
F-S
F
F
F
F
F-S

Sf
Sf
Ov
Cv
Bu
Mg
Cv
Ov
Mg

Rg
Rg
Rt
Rsp
Rg
Rsp
Rsp
Rsp
Rsp

B
B
F-S
F
F
F
F
F-S
F

Hendrickson, G. 0. (1938)
Hickie, P. (1940)
Hosley, N. W., and R. K. Ziebarth (1935)
Johnson, B. C., and A. M. Pearson (1948)
Korschgen, L. J. (1966)
Korschgen, L. J. (1967)
Latham, R. M., and C. R. Studholme (1952)
Lay, D. W. (1965)
Mosby, H. S., and C. 0. Handley (1943)

Sumac
Species

Part
Eaten

Rsp
Rsp
Rt,Rg
Rg
Rsp

Season
of
Use

Su
W,W
W
NS
Sp,F,W
W
w

NS

Preference
or
Importance

3
5,1
2
5
2
3
1 '
4

w
w

3
3

NS
NS

5
3-4

W

5
1
2
3
3
4
6
4
3

w
w
w
w

W,Sp
w

NS
w

State

Ala,
Mich.
Me.
Kans.
N.Y.
Wash.
Wise.
Mo.
Io.
Io.
Mass.
Ala.,
Miss,,
La.
Io.
Mich.
Mass.
Ala.
Mo.
NS
Penn.
Tex.
Va.

Table 8 (Cont'd)

Game
Species

Sumac
Species

Part
Eaten

Murie, A. (1946)
Murphy, D. A. (1968)
Nestler, R. B., and W. W. Bailey (1944)
Packard, R. L. (1956)
Parmalee, P. W, (1953)
Pearson, A. M. (1943)
Riegel, A. (1942)
Stoddard, H. L. (1931)

Mg
Ov
Cv
Sn
Cv
Ov
Sf
Cv

Rg
Rg
Rg
Rg
Rsp
Rsp
Rg
Rsp

F
F-S
F
B
F
F-S
B
F

Swank, W. G. (1944)
Trippensee, R. E. (1938)

Pc
Sf

Rt
Rg.Rt

F
B

Author(s)

Game Species:
Ov: White-tailed deer
Bu: Ruffed grouse
Cv: Bobwhite quail
Pc: Ringed-neck pheasant
Lc: California quail
Pp: Sharp-tailed grouse
Mg: Wild turkey
Sf: Cottontail rabbit
Part Eaten:
Sumac Species:
F: Fruit
Rg: Rhus glabra
S: Stem
Rt: Rhus typhina
B: Bark
Rsp: Rhus species not specified
Preference or Importance:
1: Preferred
2: Important
3: Moderate
4: Low
5: Mentioned, not ranked
6: Emergency

Sn:
Tc:

Season
of
Use

Preference
or
Importance

State

5
1
3
6
5
3
6
3

Ariz.
Mo.
NS
Kans.
Tex.
Ala.
Kans.
NS

3-4
5

Mich.
Mich.

W
W

w
w.sp
w
w
w
W,Sp,Su
aA

w
w

Fox squirrel
Prairie chicken

Season
W:
Sp:
Su:
A:
NS:

of Use:
Winter
Spring
Summer
Autumn
Not specified

w

27
(Goodrum and Reid, 1962), but of prime importance in the northeast.

It

is reported to be a highly preferred and important winter browse in
Maine, Massachusetts, Missouri and Wisconsin (Banasiak, 1961; Dahlberg
and Guettinger, 1956; Hosley and Ziebarth, 1935; Murphy, 1968).

Availability
About one percent of the study area, or 29 acres, was covered by
sumac.

Virtually all of the sumac present was in forest openings, and

was short enough that it was within the reach of deer.

Snow depth was

the major factor that prevented the deer from browsing the plant.

First,

there were no large deer yards on the study area, and thus few deer were
present during severe winter conditions.
coveredsome of the stems; a

Second, snow occasionally

14-inch snowfall, a typical

depth inthis area, covered about one half of

the stems.

winter snow
Sumac fruits,

however, were rarely covered by snow because they are terminal and
usually present on older, taller plants.
The relative availability of several species of deer browse in
forest openings during deep snow conditions was measured in February
1968.

The results of that survey, summarized in Table 9, demonstrated

Table 9.

Frequency and density of woody plants emerging from a 14-inch
snow cover in forest openings, as determined by 50 randomly
selected 60 square foot plots (6 ' x 1 0 ') sampled on the study
area in the winter of 1967-68

Species

Rhus typhina>glabra
P-inus ves'inoca
Rubus atleghen-iensis
Prunus serotina
Prunus virginzana
Salix humtlis
Querous rubra

Frequency
(Percent)

Stems/Acre

32
20
6
3
1
1
1

1587
315
86
43
71
43
14

28
that although many of its stems were covered by snow, sumac was the most
plentiful browse species in openings on the study area during typical
winter conditions.

Use

The use of sumac stems and fruits by deer was monitored on the
study area from July 1967 until June 1969.

The heaviest browsing

occurred in winter and thus that season received the most attention
during the study.

The winter population (post harvest) of deer in the

area was estimated to be 21 deer per section in 1967-68 and 22 deer
per section in 1968-69 (personal communication, George Irvine, Wildlife
Biologist, U. S. Forest Service, Cadillac).

The population of deer

undoubtedly changed throughout the winter, and these estimates are
intended only as convenient reference points.

Deer tracks and pellet

groups were abundant on the study area during the relatively mild winter
of 1967-68, and indicated that the area was being used by deer for the
entire season.

In contrast, very few deer signs were noted on the study

area in the winter of 1968-69.

This was probably caused by severe

weather in January of that winter, which forced deer to seek the shel­
ter of cedar swamps away from the study area.

Perhaps a contributing

factor to reduced deer use of the area was the establishment of a snow­
mobile trail along Wagon Wheel Road, and the power line right-of-way
(Figure 1).

This trail was heavily used by recreational snowmobiles

during the winter weekends.
Length of stems and fruits browsed:

Two hundred sumac stems, 50

in each of 4 clones on the study area, were measured and tagged in
August of 1968 (Figure 1).

Those tagged stems that had been browsed

were remeasured in April 1969.

A total of 100 of the 200 stems had

29
been browsed and the average length browsed was 2.4 Inches.

Eighty-two

percent of the browsed parts of the stems were between one and three
inches in length and the greatest length browsed was ten inches.

There

was no apparent relationship between the length of the stem and the
length of the browse.
Fifty fruits were tagged in order to determine how much of the stem
was eaten when a fruit was browsed.

Examination of these tagged speci­

mens after browsing showed that the fruit stem is browsed only down to
the proximal end of the fruit.

The stem easily breaks at this point and

observations of deer eating sumac indicated that they grasp the fruit
in their mouths and break it loose with a snap of the head.

Use of stems:

The seasonal use of sumac stems was measured in

eight permanent plots (501 x 100') by periodically counting the number
of stems browsed in those plots throughout the year.

When a browsed

stem was counted, it was clipped at a 50- to 60-degree angle to prevent
a recount.

Three of these plots are located on the study area and a

fourth a few hundred feet north of the study area (Figure 1).

The four

other plots are located three miles ESE of the study area in sections
13 and 24 of T21N-R12W.

The four latter plots were established before

the study area had been narrowed to its final size.
The results of the plot counts are shown in Figure 7.

Some brows­

ing occurred in the spring and early summer of 1967, but very little in
the corresponding seasons of 1968.

In the winter of 1967-68, the most

intense browsing occurred between December and March, the peak months
being January and February.

Deep snow prevented counts in January and

February of 1969, but a lower total for that winter indicated that fewer
stems were browsed in that period.

This lower total is probably a result

212B

2031

1100-

1043

1000-

700 -

600 -

550

547

OF

STEMS

BROWSED

900-

NUMBER

500 -

438

400 300 208

200

100

-

68

44

MAYJU L

ADC

71

32

17

15

I

I

I

SEP

OCT

NOV

DEC

I
JAN

FEB

MAR

APR

I
MAY

I
JON

26
I
JU L

I
AUGSEP

I
OCTNOV

I
DECAPR

MONTHS

Figure 7. The monthly consumption by deer of sumac stems on the browse plots from
May 1967 through April 1969.

31
of the deep snow, which covered many plants, and hindered the movements
of deer.
An estimate of the percentage of sumac stems browsed on the study
area from May 1967 through April 1968 was determined by tallying the
percent of stems browsed in a random sample of 100 plots (3 1 x 1 0 ') .
Sixty-three percent of all sumac stems examined had been browsed.

This

figure is comparable to the average of the percentages of browsed stems
in the four permanent browse plots on the study area, 61 percent, and
somewhat higher than the percent of stems browsed on the four plots east
of the study area, 42 percent.
Early in May 1969, an estimate of the stems browsed on the study
area during the previous year was made by tallying the percent of
browsed stems in 100 randomly chosen semicircles four feet in diameter.
Forty-eight percent of the stems were browsed, a result which compares
closely with the number of stems browsed in the browse length experi­
ment, 50 percent, but considerably higher than the number of stems
browsed on the permanent browse plots, 27 percent.

Assuming that 50

percent of all stems were browsed, and using the productivity data
for browse stems derived from Table 4, it is estimated that the deer
consumed about 200 pounds (oven-dry weight) of sumac stems on the study
area during the winter of 1968-69.

This averages out to consumption of

about seven pounds of stems per acre of sumac.

Use of fruits:

In September 1967, three groups of 100 fruits each

were tagged on the study area (Figure 1).
samples were below six feet in height.

The fruits in each of these

In another sample 100 fruits

exceeding seven feet in height were tagged on an area two miles east of
the study area.

Additionally, 70 fruits were counted inside a deer-proof

32
fence on the study area.

The rate of disappearance of the fruits that

were below six feet in height is shown in Figure 8 .

It appears that

the

fruits were lightly browsed

in September and October, with most of

the

fruit eaten during November

and December.The fruits

that were

above seven feet in height and thus out of the reach of deer, disappeared
slowly, with 80 percent remaining at the end of April and 53 percent
remaining by mid-June.

Sixty-six of the 70 fruits In the enclosure

remained at the end of the winter.

These experiments were repeated in

1968-69 with very similar results (Figure 8 ).
According to the estimate of fruit production in 1968 (Table 4),
nearly 2,300 pounds
the

(oven-dry weight) of sumac

study area from November to

fruit were browsed on

mid-January ofthe winter

of 1968-69.

This averages out to a consumption of about 80 pounds of fruit per acre
of sumac.

The impressive statistic here is that all of the fruits

available to deer were browsed, suggesting that the consumption of sumac
fruits would have been even higher if more fruits had been available.

Summary
A survey of the literature indicated that smooth and staghorn sumac
are important wildlife food plants throughout much of their ranges in
North America.

The present study demonstrated that sumac was heavily

utilized as a winter food by deer In the Manistee National Forest in
Michigan,
(1)

The major findings of this study are outlined below:
Sumac was the most abundant browse species available in

forest openings during typical winter conditions on the study areas.
(2)

The average length of stem browsed was 2.4 inches and In 82

percent of the cases the length of stern browsed was between one and
three inches.

33

70H
60 H

50H

PERCENT

OF

FRUIT

REM AINING

100 H

1 9 6 7 -6 B

AUG

SEP

OCT

NOV

DEC

JA N

MONTHS

Figure 8 . Rate of disappearance of sumac fruit susceptible to
deer browsing, on the study area, in the autumn and winter of
1967-68 and 1968-69.

34
(3)

In the winter of 1968-69, approximately half of the sumac

stems on the study area were browsed.

This totaled about 200 pounds

(oven-dry weight) of sumac stems, or about seven pounds of stems browsed
per acre of sumac.
(4)

When a fruit was browsed, the entire fruit was eaten.

(5)

All of the sumac fruit on the study area was browsed in the

winter of 1968-69.

This totaled about 2,300 pounds of sumac fruit

browsed on the study area, or about 80 pounds browsed per acre of sumac.

THE PROXIMATE ANALYSIS AND APPARENT DIGESTIBILITY OF SUMAC

As demonstrated in Chapter 3, sumac may comprise a significant
portion of the winter diet of deer in Michigan.

In an attempt to cor­

relate the use of this browse with Its nutritional value,

the stems and

fruits of smooth and staghorn sumac, and the hybrid sumac, were analyzed
for nutrient composition.

Further, apparent digestibility trials were

conducted with the fruit of smooth sumac.

Methods and Materials

Proximate analyses were conducted on a total of 19 samples:
stem samples, 3 fruit samples, 1 orts sample and 2 fecal samples.

13
All

samples were weighed fresh, air dried, and ground in a Wiley Mill with
a #40 mesh screen.

Subsequently, their percentages of crude protein,

ether extract, ash and dry matter were determined by A.O.A.C. procedures
(Horwitz, 1960).

Percentages of the cell wall constituents, cellulose,

hemicellulose, and lignln, were determined by processes outlined by
Van Soest (1963) and Van Soest and Wine (1967).

These methods differ

from standard fiber determinations (Horwitz, 1960), but are believed to
provide a more accurate determination of the fibrous and soluble carbo­
hydrate fractions (Van Soest, 1963, 1967; Fonnesbeck, 1968, 1969).
Calcium and magnesium content was determined by means of a JarrellAsh Atomic Absorption Spectrophotometer, and phosphorus content was
determined by means of a Beckman Spectrophotometer.
Calorimeter was used to obtain gross energy values.
35

A Parr Isothermal

36
The letter designations (A through P) from Table 10 will serve to
denote each sample.

Samples A through C are each composed of stems

from six clones of their respective species, and each clone is repre­
sented equally by weight.

Sample D is represented by only two clones.

Samples E and F were taken from the root sprouts of adjacent clones

Table 10.

Desig­
nation

Location of sumac clones used in analyses

Date Col­
lected (1969)

Species

Part

County (number of clones)

A

R. typhina

stem

Clinton (2); Ingham (2);
Shiawassee (2)

Feb. 25

B

R. glabra

stem

Clinton (2); Washtenaw (2);
Ingham (1); Shiawassee (1)

Feb. 25

C

R. t>g*

stem

Wexford (6 )

Feb. 28

D

R. t>g

s tern

Ingham (2)

Feb. 26

E

R. glabra

s tern

Ingham (1)

Feb. 26

F

R. typh-ina

stem

Ingham (1)

Feb . 26

G-M

R. typhina

stem

Clinton (1)

Feb. 25

N

R. typhina

fruit

Ingham (3); Clinton (3)

Feb . 25

0

R. glabra

fruit

Washtenaw (6 )

Feb. 1-3

P

R. t>g

fruit

Wexford (6 )

Feb. 26

*R. t>g = Rhus typhina >glabra

that had been clearcut prior to the last growing season.

Only the first

2-1/2 inches of each stem was utilized in samples A through F, as this
length corresponds closely to the average length of stem browsed by
deer.

Samples G through M are consecutive one-inch segments of 50 stems

37
taken from one clone with sample G representing the first inch and
sample M the seventh.
The fruit samples N and P represent composites from six clones of
their respective species.

Sample 0 is a subsample of the fruit used in

the apparent digestibility trials.
All samples were collected in February 1969, and represent the pre­
vious year's growth.

The stems and fruits of all samples were randomly

collected within each clone, and all diseased or otherwise damaged
specimens were discarded.

Upon collection all stem samples were placed

in tared plastic bags, frozen, and their fresh weight was determined.
The fruit samples, which were air dried naturally in the field, were
placed in cloth bags upon collection.

The samples used in the digesti­

bility trials were stored in cloth bags inside an unheated barn until
they were used.
Apparent digestibility trials, in which smooth sumac fruit was fed
to deer, were conducted at the Houghton Lake Wildlife Research Station,
Houghton Lake, Michigan.

This study began on 28 January, 1969, and

originally employed six deer.

The deer, three 4-1/2-year-old does and

three 1 -1 /2 -year-old bucks, were born in captivity, and fed a commercial
feed prior to the experiment.
The apparent digestibility study was divided into three consecutive
phases.

Phase I was a seven-day adjustment period, during which the

deer were presented both sumac and the commercial feed.

The commercial

feed was gradually withdrawn until, at the end of seven days, the deer
were being fed only sumac.

Phase II, which followed immediately, lasted

17 days, during which the deer were fed only sumac.

During Phases I and

II the deer were segregated by sex and kept in open, outside pens.

The

38
average minimum and maximum temperatures during these periods were
2 and 29° F, respectively.
At the end of Phase II it was determined, by visual inspection,
that the three does and one of the bucks had suffered severe weight
loss and they were withdrawn from the experiment.

The two remaining

bucks were deemed healthy enough to continue on the experiment, although
they also had suffered noticeable weight loss.
In Phase III the two bucks were placed in metabolism cages within
a heated barn, for a period of 14 days.

During this period they were

fed sumac and water ad X'Lb'Lbum> and were subjected to a minimum of dis­
turbance.

The temperature within the barn varied from 40 to 54° F.

The deer were fed and watered each morning, at which time the feces
and urine which had accumulated over the past 24 hours were removed and
weighed.

The food and water were weighed in and weighed out to determine

total consumption.

Uneaten food from the previous day (orts) was saved

and later analysed to determine if the deer were selectively eating the
fruit.

Aliquots of the urine and feces for analysis were taken daily

during the last seven days of Phase III.
H 2 SO 4 was added to the urine sample,

Ten milliliters of 0.1 N

thus lowering its pH, and reducing

the loss of ammonia nitrogen.
The metabolism cages measured 4' x 4 1 x 4', and were entirely
wooden except for a metal grill floor,
passed.

through which feces and urine

Fecal material, after passing through the grill, was caught by

a wire screen.

Urine passed through both the grill and screen and was

intercepted by a laquered wooden surface which funneled it into a col­
lection pan.

Panels in the roof of the cage permitted air circulation

and the entrance of light.

39
Results
The results of the proximate analyses appear in Table 11.

The com­

posited stem samples, A through D, appear fairly similar, with no major
differences occurring between the samples.

The one-year-old stems from

the clearcut area, E and F, differ principally in their percentages of
hemicellulose.

Samples E and F were higher in several categories of

nutrients than samples A through D.

However, this increase in nutrients

was probably a reflection of the higher percentage of dry matter in
those samples.
In stem samples G through M there was an obvious decrease proximally in the soluble fraction.

Correspondingly, the fibrous fraction

increased proximally except for lignin, which did not demonstrate a
trend.

The percentage of ash tended to decrease proximally, as did the

percentage of calcium, but no trends were discernible for phosphorus
and magnesium.

The values for gross energy did not vary significantly

among the stem segments.
The staghorn and smooth sumac fruit samples, N and 0, appeared
fairly similar, differing most notably in their cellulose and soluble
carbohydrate fractions.

The hybrid fruit, P, had a nutrient makeup

which varied somewhat from the other fruits, the greatest difference
occurring in the percentage of ether extract.

There was little differ­

ence in the values of gross energy between the fruits.

The orts, Q,

were not considered to be sufficiently different from the presented
food, 0 , to correct for selection on the part of the deer during the
apparent digestibility trials.
The results of the apparent digestibility trials appear in Table
12.

Both deer consumed similar amounts of food and water.

The apparent

digestibility of cellulose, soluble carbohydrate and ether extract was

Table 11.

Proximate analyses of sumac stems and fruit, expressed on a fresh weight basis

A
Staghorn
Sumac
Stem

B
Smooth
Sumac
Stem

C
Hybrid
Sumac
Stem

D
Hybrid
Sumac
Stem

E
Smooth
Sumac
Stem

F
Staghorn
Sumac
Stem

G
Staghorn
Sumac
1st Inch

H
Staghorn
Sumac
2nd Inch

56.85

59.29

55.46

54.62

62.80

64.00

55.52

55.50

Fibrous fraction (Percent)
Cell wall constituents
Cellulose
Hemicellulose
Lignin

22.98
11.80
4.60
6.58

25.22
12.57
5.24
7.41

23.68
12.06
5.26
6.36

23.06
11.03
3.03
9.00

24.12
13.72
2.55
7.85

25.90
13.84
3.50
8.56

22.76
11.05 .
4.43
7.28

26.50
13.78
5.03
7.69

Soluble fraction (Percent)
Cellular contents
Soluble carbohydrates
Protein+
Ether extract
Ash
Calcium
Phosphorus
Magnesium

33.87
20.61
3.85
6.22
3.10
0.86
0.12
0.16

34.07
21.01
3.45
6.54
3.08
1.13

31.78
19.06
3.90
6.08
2.74
*

0.17

0.14
0.15

31.56
20.04
4.04
4.82
2.66
0.90
0.09
0.15

38.68
23.38
4.26
7.46
3.58
1.21
0.09
0.17

38.10
23.10
4.38
6.93
3.60
1.04
0.11
0.24

32.76
19.81
3.80
5.71
3.44
1.55
0.12
0.21

29.00
17.67
3.64
5.04
2.65
1.05
0.14
0.18

2.835

2.666

Dry matter (Percent)

energy (kcal/g)

2.717

2.556

3.040

2.942

2.644

2.474

Table 11 (Cont'd)

M
Staghorn
Sumac
7th Inch

N
Staghorn
Sumac
Fruit

0
Smooth
Sumac
Fruit

D
4.
Hybrid
Sumac
Fruit

Orts

55.85

55.75

94.06

94.06

94.10

93.35

30.94
17.23
6.70
6.92

32.19
17.37
7.11
7.71

32.48
18.59
6.66
7.23

49.77
18.88
15.23
15.66

53.59
21.68
14.05
17.86

47.78
17.76
16.83
13.19

51.53
22.33
12.30
16.90

24.44
14.30
3.37
4.46
2.31
0.94
0.12
0.18

23.66
14.37
3.17
4.15
1.97
0.80
0.10
0.19

23.27
13.91
3.07
4.10
2.19
0.70
0.09
0.15

44.69
22.83
5.01
13.89
2.96
0.60
0.17
*

40.49
18.10
5.24
13.80
3.35
0.85
0.13
0.23

46.28
16.65
6.42
20.65
2.56
ft
0.15

41.82
18.72
4.79
15.33
2.98
0.92
0.20
0.24

I
Staghorn
Sumac
3rd Inch

J
Staghorn
Sumac
4th Inch

K
Staghorn
Sumac
5th Inch

55.37

55.87

55.38

Fibrous fraction (Percent)
Cell wall constituents
Cellulose
Hemicellulose
Lignin

27.91
14.68
5.54
7.69

30.27
15.28
6.84
8.15

Soluble fraction (Percent)
Cellular contents
Soluble carbohydrates
Protein+
Ether extract
Ash
Calcium
Phosphorus
Magnesium

27.46
16.81
3.63
4.64
2.38
0.94
0.13
0.16

25.60
15.24
3.41
4.45
2.50
0.96
0.11
0.17

Dry matter (Percent)

Gross energy (kcal/g)

+N x 6.25
*Data missing

2.497

2.565

2.446

L
S taghorn
Sumac
6th Inch

2.441

2.637

5.002

4.707

•.I-

4.809

Q

4.425

42
Table 12.

Sumac browse and water Intake, apparent digestibility and
apparently digestible energy during the last seven days of
Phase III

Average daily intake (Kg.)
Sumac fruit*
Water
Apparent digestibility (Percent)
Dry matter
Cell wall constituents
Cellulose
Hemicellulose
Lignin
Cellular contents
Soluble carbohydrates
Crude protein
Ether extract
Gross energy
Apparently digestible energy
intake per day (kcal)***

Deer 1

Deer 2

Mean +
S tandard
Error

0,563
0.951

0.521
.
1 .122

0.542+0.021
1.036+0.135

52.6
43.3
47.8
83.4
19.9

39.7
29.0
50.9
36.3
-1.5

46. 1+ 6 .4
36. 1+7.1
49. 3+1.5
49. 8+13.5
9. 2+ 1 0 .7

68.7
24.2
85.5

71.2
-56.5
81.9

69. 9+1.2
-16. 1+40.4
83. 7±1 .8

50.8

37.5

1430

977

44, 1+ 6.6

1204+227

*Oven-dry weight
**NX6.25
***Gross energy intake X apparent digestibility of gross energy

similar for the two animals, but large discrepancies occurred between
the deer in the digestibility of protein and hemicellulose.

Lignin was

not considered to be a digestible fraction of a deer's diet.
The apparent digestibility data obtained for gross energy were
further refined by determining the amount of energy lost in the urine
and through methane production (Table 13).

Methane production was

estimated by the procedure outlined by Blaxter and Clapperton (1965).
This refined gross energy determination was termed apparently metabo­
lizable energy.

Figures for both apparently metabolizable and

A3
Table 13.

Urine and methane output, nitrogen balance, and apparently
metabolizable energy during the last seven days of Phase III

Average daily output
Urine (ml.)
Nitrogen (gm.)
Energy (lccal)
Hethane*
Energy (cal)
JL JL

Nitrogen balance

,

,

(gm.)

Apparently metabolizable
energy intake per day (kcal)***

Mean +
Standard Error

Deer 1

Deer 2

537
11.1
16A

898
17.1
202

717+181
14.1+3.0
183+19

192

156

174+18

-9.8

-19.8

- 1 A .8+5.0

1073

501

787+286

*Assumes maintenance level of feeding (Blaxter and Clapperton, 1965).
**Nitrogen Intake - nitrogen output in feces and urine.
***Gross energy intake - gross energy output in feces, urine
and methane.

apparently digestible energy were given because the latter, though less
precise, is necessary for comparisons of work done by other investigators.
The nitrogen balance of the deer during the collection period was
determined by comparing the total Intake and output of nitrogen (Table
13) .

Both deer were in negative nitrogen balance, indicating that pro­

tein catabolism was proceeding more rapidly than protein synthesis.
From the beginning of Phase I until the end of Phase III, a period
of A5 days, deer 1 and 2 lost 28.2 percent and 31.8 percent of their
body weights, respectively.

The weight losses during the lA-day period

in the metabolism cages were 16.A percent for both deer (Table 14).

44
Table 14.

Weight losses during Phases I-IITW of the digestibility
study

Phases 1-.II
Beginning Final Percent
Weight
Weight Weight
Deer
(leg.)
(kg-) Loss

Phase III
Beginning Final Percent
Weight
Weight Weight
(kg.) Loss
(kg.)

Phases
I-III
Percent Total
Weight Loss

I

64.4

55.3

14.1

55.3

46.2

16.4

28.3

2

61.2

49.9

18.4

49.9

41.7

16.4

31.9

"Phases I and II: January 28 to February 20.
Phase III: February 21 to March 7.

Discussion
The results obtained from the proximate analyses in this study were
similar to analyses of smooth sumac fruits collected in Michigan, V i r ­
ginia, and Washington by King and McClure (1944), and of smooth sumac
stems collected in Missouri by Murphy (1968).
The reader is cautioned that the values presented

in this paper

represent only a relatively small number of vigorous clones.

Time limi­

tations prevented analysis of nutritional differences among the factors
of site, season, age, vigor, genotype, or position

on the plant.

Nutri­

tional differences in other species of deer browse

due to such variables

have been shown by Bailey (1967), Bissel and Strong (1955), Broadfoot
and Farmer (1969), Einarsen (1946), Forbes

(1941), Helmers (1940)

and Swift (1948).
Proximate analyses are of importance from the standpoint of deter­
mining potential nutrient concentration.

Such data are limited in

determining the nutritional value of a browse unless accompanied by
information on the preference, consumption, availability and digesti­
bility of that browse.

In the absence of digestibility data on smooth

45
sumac stems and staghorn sumac stems and fruits, it is perhaps most
instructive to compare their proximate analyses with those of deer
browse from northern white cedar (Thuja oaaidentaZ'is) , big tooth aspen

(Populus grandidentata ) , and jack pine (Pinas banksiana ).
of these browses and the sumac browses appear in Table 15.

Table 15.

The analyses
The data are

A comparison of the proximate analyses of several winter
browses.
(All analyses except dry matter are expressed on
an oven-dry weight basis.)

Percent
Dry
Matter

N. white cedar, sprays*
Big tooth aspen, stems**
Jack pine, boughs***
Smooth sumac, s terns
Staghorn sumac, stems
Hybrid sumac, stems
Smooth sumac, fruits
Staghorn sumac, fruits
Hybrid sumac, fruits

46.1
51.7
46.5
59.3
56.8
55.5
94.1
94.5
94.1

Percent
Crude
Protein

Percent
Ether
Extract

Percent
Ash

Gross
Energy
(kcal/g)

7.2
9.7
8.2
5.9
6.7
7.0
5.5
5.3
6,8

9.5
6.8
9.0
10.9
10.9
10.9
14.6
14.6
21.9

4.3
3.7
2,6
5.2
5.6
4.9
3.5
3.2
2.7

5.14
5.01
5.36
4.77
4.78
4.80
5.00
5.28
5.11

*Ullrey et at. , 1968
**Ullrey et at. , 1964
***Ullrey et al. , 1967

expressed on an oven-dry basis to eliminate differences due to water
content.

Because of differences in analytical techniques among research­

ers, only the dry matter, protein, ether extract and gross energy frac­
tions can be compared.

The stems and fruits of all sumac species were

lower in protein and higher in ether extract than the other browses.
The ether extract of the hybrid fruit was especially high.

The ash con­

tent of the sumac stems was comparatively high, while that of the fruits

46
was intermediate.

None of the browses differed greatly in gross energy

content.
The stem segments (G through M) demonstrated a progressive decrease
in the soluble fraction, and a corresponding increase in the fibrous
fraction, proximally.

Bailey (1967) has found a similar occurrence in

the protein fraction of V-ibuvnum sp. stems.

Short (1963, 1966) has

demonstrated that deer derive the most benefit from the soluble fraction
of a food, and that the value of a browse is inversely related to its
cellulose content.

The distal increase in fiber of a sumac stem may

partially explain why only the first few inches are browsed.
The intake and apparent digestibility of cedar and sumar are com­
pared in Table 16.

Although the consumption of cedar and sumac differ

greatly when compared on a fresh weight basis,

they are essentially the

same on a dry weight basis and provide similar amounts of apparently
digestible energy.

These data also indicate that the protein fraction

of cedar is more available to deer than that of sumac, while the reverse
appears to be true in regard to the availability of ether extract.
Ullrey et at.

(1969) have estimated that the apparently digestible

energy requirement for the winter maintenance of deer in Michigan is
about 160 kcal/kg W

1 cA

/day.

Thus a deer of 138 pounds,

the average

beginning weight of the deer in this study, would require 3565 kcal/
day of apparently digestible energy.

Assuming a gross energy digesti­

bility of 44 percent (Table 12), sumac fruit produced about 1000 kcal
of apparently digestible energy per pound (oven-dry weight).

* W 75 refers to metabolic weight (Kleiber, 1961).

Table 16.

A comparison of cedar sprays and sumac fruits in terms of intake, apparent digestibility,
apparently digestible energy, and weight loss

No.
of
Deer

N. white
cedar,
*
sprays

Smooth
sumac,
fruit

24

2

Browse Intake per
Deer per Day
Fresh
Oven Dry
Weight
Weight
(kg.)
(kg.)

Apparent Digestibility
Percent
Percent
Percent Percent
Gross
Crude
Ether
Dry
Protein
Extract Energy
Matter

Apparently
Digestible
Energy/
Deer/Day
(kcal)

Percent
Weight Loss

1.240+**
0.108

0.572+
0.049

44+4

14+4

47+5

39+3

1140

12.1+1.4

0.578+
0.021

0.542+
0.021

46+6

-16+40

84+2

44+7

1204

16.4+0.0

*Ullrey et at., 1968
**Mean + standard error

48
The deer in this study consumed enough food to supply about onethird of their maintenance energy requirements and, after 31 days on
the pure sumac diet, they had lost about 30 percent of their original
body weight.

Additionally, both deer were in negative nitrogen balance.

Continued protein and calorie malnutrition would have soon resulted in
death from starvation.
Several authors have commented on the voluntary restriction of
food intake by deer during the winter, resulting in considerable weight
loss (French et at . , 1936; McEwen et a t ., 1957; Silver et at . , 1969;
Smith, 1950).

Silver et at . (1969) have associated voluntary reduction

in food intake during the winter with a reduced metabolic rate.

They

suggest that this response is a physiological and behavioral reaction
of deer forced to exist in areas where food is limited during the winter.
Ullrey et a t . (1964, 1967, 1968) have consistently encountered under­
nourishment, primarily due to inadequate food intake, in deer fed a
diet consisting of only a single browse species.

This may indicate that

deer do not adapt easily to a monotypic diet.
The present study was not designed to explain the inadequate food
intake, and no further attention will be given to that subject.

The

data from the digestibility trials do not lend themselves to statisti­
cal analysis because only two of the original six deer finished the
experiment.

However, they do seem to indicate that sumac fruit is a

high energy food source, but offers little available protein.

Summary
The stems and fruits of smooth, staghorn, and the hybrid sumac were
analyzed for nutrient composition, and the apparent digestibility of
the fruit of smooth sumac was determined.
the study are outlined below:

The principal findings of

49
(1)

There appeared to be little interspecific difference between

the nutritional composition of the sumac stems and fruits.
(2)

Sumac stems and fruits were low in crude protein, high in

ether extract, and similar in gross energy as compared, on an oven-dry
weight basis,

to three other Michigan deer browses:

northern white

cedar, jack pine, and big tooth aspen.
(3)

Proximate analyses of consecutive one-inch stem segments

demonstrated that the soluble nutrient fraction decreased, while the
fibrous fraction increased, proximally.
(4)

Six deer were placed on a diet consisting only of smooth

sumac fruit.

Two of these deer had adjusted adequately to the sumac

diet after three weeks, and these animals were used in the subsequent
apparent digestibility trials.
(5)

The two deer had similar apparent digestibility percentages

for cellulose, 49.3+1.5 (mean + standard error); soluble carbohydrates,
69.9+1.2; and ether extract, 83.7+1.8.

They differed somewhat in the

apparent digestibility of gross energy, 44.1+6.6, and differed greatly
in the apparent digestibility of crude protein, -16.1+40.4.
(6)

Both deer voluntarily restricted their intake of sumac fruit

to about one-third of that amount required to maintain their body
weight.

Accordingly, both deer were in negative nitrogen balance and

both suffered severe weight loss.
(7)

The digestibility data indicated that sumac fruits were a

good energy source, but a poor source of protein, compared, on a dry
weight basis, to sprays of northern white cedar.

ESTABLISHMENT AND PROPAGATION OF THE PLANT

Methods of establishing sumac from both seed and transplants, and
of rejuvenating established plants to render them more useful to wild­
life, are described.

All of these studies were conducted in Clinton,

Ingham, and Shiawassee Counties and were concerned primarily with ele­
mentary nursery practices.

Collection and Germination of Seeds
Sumac seeds are oval, smooth, 2 to 3 mm. long, and have an extremely
hard seed coat.

They exhibit mechanical dormancy, and acid scarifica­

tion is the most commonly recommended procedure employed to prepare the
seeds for germination (Boyd, 1943a; Heit, 1967a; Rrefting and Roe, 1949;
Lovell, 1964).

The seeds do not require special storage procedures

either before or after scarification, and may be kept in a sealed con­
tainer for several years without loss of viability (Heit, 1967b).

Methods:
October.

Sumac seeds were collected by harvesting ripe fruit in

The seeds were removed from their leathery pericarp by drying

them for three days at 110° F., placing them in a cloth bag (one-half
pound of fruit in a 10" x 16" seed b a g ) , and vigorously pounding the
bag against a hard surface for a period of two minutes.

The contents

of the bag were then placed in a tray of water and stirred.

The viable

seeds, which had a greater density than water, sank to the bottom of
the tray, while the fruit debris floated and was easily removed.
50

The

51
water was then decanted and the seeds were retrieved, air dried, and
stored in a capped glass bottle.

This process was repeated many times

during the course of the study with all three sumac types.
In order todetermine the optimum length
cation, 50

of time for acid scarifi­

gramsof Rhus typhincp glabra seeds were placed in 500 ml.

of

concentrated sulfuric acid and stirred constantly with a magnetic
stirrer.

Aliquots of the seeds were removed from the acid at the fol­

lowing intervals:
and 480 minutes.

10, 20, 30, 40, 60, 90, 120, 180, 240, 300, 360, 420
Upon removal from the acid the seeds were washed

thoroughly in water and the burned portion of the seed coat was removed
by gently rubbing the seeds over a fine wire screen.

One hundred seeds

from each sample were placed between two filter papers (Whatman ill) in
a sterile petri dish.

The seeds were kept under conditions of constant

light and temperature (73-76° F ) , and sufficient distilled water was
added to the filter paper to make it moist to the touch.

Additional

water was added when necessary, to maintain the moist condition.

Each

day for 42 days the dishes were opened and seeds which had germinated
were removed.

A seed was considered to have germinated when root hairs

appeared on the root tip.

Results and discussion:

There was considerable inter- and intra­

specific variation in the size and weight of the seeds obtained.

The

seeds of Rhus glabra were significantly heavier than those of Rhus

typhina (P< .05),*
overlapped

and the size of the seeds of Rhus typh'iria> glabra

those of the parent species (Table 17).

*Mann-Whitney Test, Siegel (1956), p. 312.

52
Table 17.

Weight of sumac seeds per fruit and number of seeds per pound

Pounds of viable
seeds per 100
pounds fruit

Clones
Sampled

Species

Rhus glabra
Rhus typhina
Rhus typhina>glabra

4
4
4

Number of clean
viable seeds per
pound

13.7+5.1
14.4+4.5
22.9+5.2

46,400+3,200*
60,200+2,500
51,900+3,900

^Standard error of the mean

The results of the scarification-germination experiment are shown
in Figure 9.

Only two percent of the unscarified controls, and two per­

cent of the seeds scarified for 10 minutes, germinated.

The optimum

time of scarification was 180 minutes, beyond which time the acid
apparently penetrated the seed coat and damaged the seeds.

None of the

seeds scarified for 420 minutes germinated, and those scarified for 480
minutes were nearly dissolved.

Seventy percent of all of the seeds

that germinated did so within the first four days, and 93 percent of
the total had germinated within seven days (Figure 10).
All of the seeds used in subsequent planting trials were scarified
for 120 minutes, slightly less than the optimum time, to avoid possible
damage to the embryo that may have gone undetected in the germination
trials.

Thus the optimum expected germination in all planting trials

is only 88 percent (Figure 9).

Propagation from Seed
Sumac reproduces primarily from root sprouts, but natural original
establishment must depend upon seeding.

Several authors have demonstrated

that seed germination in several sumac species, and hence reproduction

100-

90 -

-

PERCENT

OF

SEEDS

FERM IN ATED

80

30 -

10

-

90

C
MINUTES

120

I N S U L F U R IC A C I D

180

240

300

360

(CONC)

Figure 9, Percentage of sumac seeds germinating after various time intervals in
concentrated sulfuric acid.

420

54

200-

1 5 0 -

125-

100

NUMBER

OF

SEEDS

GERM IN ATING

175-

75-

5 0 -

2 5 -

1

2

3

4

5

6

7

8

9

10

11

12

DAYS

Figure 10.
ac i d .

Rate of germination of seeds scarified in sulfuric

from seed, is greatly enhanced by fire (Lovell, 1964; Stone and Juhren,
1951; Went et a t ., 1952; Wright, 1931).

Additionally, sumac seed germi­

nation has been shown to increase following ingestion by mammals and
birds (Brown, 1947; Krefting and Roe, 1949; Swank, 1944).

It appears

that both animal ingestion and fire are natural methods of scarification
and may be important means of sumac establishment in some parts of
North America.

However, in six months of daily field observations in

sumac areas, I have noted only two seedlings growing in the wild.

Cer­

tainly, a sumac management plan in Michigan cannot depend upon natural
seeding and, therefore, this aspect of the study was approached from
the standpoint of artificial establishment from seed.

Methods:

In June 1968 three seeding techniques were tested at

the Michigan State University Tree Research Center using scarified seeds
of Rhus typhina>glabra .

In two of these procedures the seeds were broad­

cast, and in the third the seeds were planted.

Six 24 square foot

plots (6' x 4') were raked and each was then broadcast with 3,500 seeds,
approximately one seed per square inch.

Subsequently, three of the

plots were left undisturbed and three were raked by dragging a rake once
over the entire plot, thus covering many of the seeds with soil.

In

another plot (4* x 10') one hundred seeds were planted at each of the
following depths:

one inch, two inches, three inches, and four inches.

During the course of the summer the soil, a loamy sand, was kept moist
by rainfall and irrigation.
Laboratory experiments were also conducted at the Michigan State
University Plant Science Greenhouse in which scarified seeds of Rhus

typh'ina>gtdbTa were planted in a sandy loam soil at depths of 0.16
inches, 0.33 inches, 0.66 inches, and 1.00 inch.

Fifty-seven seeds

56

were planted at each depth.

The soil in the greenhouse trays was kept

moist, and lightly dusted with the fungicide Captan.

Results and discussion:

The results of the seeding experiment at

the Tree Research Center are shown in Table 18.

These data strongly

suggest that the seeds need to be covered with soil, but that the suc­
cess ratio drops sharply if they are planted deeper than one inch.
About 70 percent of all of the plants grown in the summer of 1968 sur­
vived the following winter.
The data from the greenhouse plantings indicated that the most
successful planting depth is between 0.16 and 0.33 inches (Figure 11).

Table 18.

The success of several seed planting trials at the Tree
Research Center

Type of Planting

Broadcast,
Broadcast,
Broadcast,
Broadcas t ,
Broadcast,
Broadcas t ,
Planted at
Planted at
Planted at
Planted at

not raked
not raked
not raked
raked
raked
raked
1"
2”
3"
4"

Number of
Seeds Used

Number of Plants
Established

3500
3500
3500
3500
3500
3500
100
100
100
100

1
10
0
373
313
407
29
4
1
0

However, a high percentage of the seeds planted at 0.66 and 1.00 inches
were successful.

Planting as deep as one inch may be advantageous under

uncontrolled field conditions, where shallow seeds would be more sus­
ceptible to drought.

57

100
90

1 /3 I n c h

2 / 3 Inch

or

SEEDS

rcK K IX A T C !

1 /ft lo c h
70-

m c iw

1 Inch
30

6

6

10

12

If

16

1?

19

Figure 11Percentage of seeds germinating when planted at
various depths in soil trays, under greenhouse conditions.

58
The assessment of the value of these results in hampered by the
absence of field data.

However, from the information obtained, it

appears that sumac could be established in the field from seed plantings.
Perhaps the most efficient method would be to broadcast seeds prior to
discing.

Transplants

Basically, two types of transplants were made.

In one, seedlings

were grown in a greenhouse and later transplanted to the Tree Research
Center.

In the second, plants grown from seed the previous year at

the Tree Research Center were transplanted a distance of only a few feet
from the site in which they were originally planted.

The experiments

were designed to determine successful methods of transplanting sumac.

Methods:

Sumac plants were grown in soil trays from seed, and in

Jiffy-7 peat pots in the Plant Science Greenhouse.

In June 1969, five

weeks after planting, 74 of the seedlings, 37 from each treatment, were
transplanted to the Tree Research Center.

The plants from the soil

trays were transplanted bare rooted, and the peat pot plants were
transplanted so that the entire pot was covered with soil.
In July 1969, 90 of the sumac plants established during the pre­
vious year were dug up and treated in one of three ways.
plants had the soil removed from their roots.

Sixty of the

Thirty of these were

replanted immediately, and 30 were replanted after their roots had been
exposed to the air (at 78° F) and sunlight for one hour.

The soil

around the roots of the remaining 30 plants was left as intact as pos­
sible when these plants were transplanted.

The roots of these plants

were balled in approximately one cubic foot of soil.

59
Results and discussion:
are shown in Table 19.

The results of the transplant experiments

All methods of transplanting were judged suc­

cessful, with the best methods being those in which fewest of the plants
lost their leaves.

Only those plants that did not lose their leaves,

or that resprouted, were considered survivors.

Table 19,

Comparison of several transplant methods attempted at the
Tree Research Center

Method of Transplant

Peat pots, planted
entire
Tray plants, planted
bare rooted
Two-year-old plants,
planted bare rooted,
roots not exposed to
drying
Two-year-old plants,
planted bare rooted,
roots exposed to
drying for 1 hour
Two-year-old plants,
planted with roots
balled in soil

Percent of Sur­
Percent of
Number Percent
vivors that lost
Survivors
of
Surviving
that did not leaves but
Plants in 2 months lose leaves
resprouted

37

81

100

37

86

100

30

86

49

38

30

77

7

70

30

97

84

13

The peat pot transplants were somewhat less successful than the
soil tray transplants, as a greater percentage of the latter survived.
However, the survivors of the peat pot treatment were significantly
larger in terms of stem length (P < .05) ,

^Unpaired t-test, Li (1964), p. 104.

and had considerably better

60
root development, in terms of root weight (P < .05)* than the survivors
of the tray transplant treatment.

Some of the peat pots were partially

unearthed by the erosive action of rainfall.

These pots, upon exposure

to the air, lost moisture rapidly and the plants within them died as
a result of the droughty conditions produced.
The most successful of the two-year-old transplant procedures was
the method in which the roots were kept intact with the soil in which
they had grown.

Most of the plants in this treatment continued growing

without losing their leaves or resprouting.

The least successful was

the method in which the roots were exposed to the air for one hour.
However, even in this procedure, most of the plants did survive, and
this may be the most practical method for large scale planting opera­
tions because it demands the fewest precautions.

Propagation by Physical Disturbance
Root severing experiments were conducted on several clones that
appeared to be dormant, in an attempt to rejuvenate them; but no responses
were noted, and it seems probable that those clones were either dead
or of such reduced vigor that response was impossible.

However, the

29 acres of sumac on the study area are evidence that sumac responds
vigorously to physical disturbance of the roots.

In that area, a vigor­

ous growth response apparently occurred when the roots of sumac in the
area were severed by a plow set at a depth of about 10 inches during
pine planting operations.

The furrows were spaced about six feet apart,

and the operations were conducted primarily in April and May.
None of the hybrid sumac plants on the study area had grown out
of the reach of deer.

However, smooth sumac and, more commonly,

^Unpaired t-test, Li (1964), p. 104.

61
staghorn sumac, often reach heights far in excess of the reach of deer,
and thus become useless as browse species.

Mowing or cutting the stems

to correct this situation is discussed below.

Methods:

Twelve areas, six each of Rhus gtahva and Rhus typhina,

in which most of the stems had grown out of the reach of deer, were
located in Ingham, Clinton and Shiawassee Counties.

In April 1969 all

plants were removed from one 400 square foot plot (20' x 20') within
each of the 12 areas.

Each plant was cut with a pruning saw at a

height of four inches from the ground.

The age of each plant was

determined, and its new growth counted and weighed.

In October 1969

the new sprouts on the area were counted, measured, and weighed.

Results and discussion:
sented in Table 20.

The results of this experiment are pre­

Each clone responded vigorously, and in each case

the browse zone was successfully lowered.

There were significant corre­

lations* (P < .05) between the number of plants cut in an area and the
number (r = .88) and weight (r = .66) of sprouts that grew back during
the following growing season.

Additionally, there were significant

correlations (P < .05) between the number (r - .67) and weight (r =
.76) of stems growing on the removed plants, and the weight of the
sprouts that grew back in the following season.

These correlations

demonstrated a positive relationship between the vigor of the clone
and its response to cutting, indicating that clonal vigor was important
in determining the results of this management technique.

Physical

disturbance by mowing thus appears to be a useful method In lowering
the height of browse in existing clones, and large scale sumac mowing

*Li (1964), p. 301.

Table 20.

Productivity of sumac before and after mowing

Number
of
Plants

Number
of
Stems

Weight*
of
Stems
(grams)

Average
Age

Weight*
of
Sprouts
(grams)

Species

Sex

Average
Height
(feet)

1

i?. typhina

F

6.5

18

72

125

7

23

1000

3.1

2

R. typhina

F

6.5

27

130

483

7

25

909

3.1

3

if. typhina

F

10.0

34

387

3328

9

102

3952

4.3

4

R. typhina

H

5.5

33

160

317

9

45

640

1.9

5

if. typhina

M

5.5

79

275

492

8

78

1012

2.1

6

if. typhina

M

7.0

135

406

1442

7

236

4477

3.8

7

if. glabra

F

5.5

52

173

490

6

64

1311

2.1

8

if. glabra

F

6.5

95

677

1598

6

112

3404

3.4

9

if. glabra

F

5.0

76

490

1980

10

73

1613

2.9

10

if. glabra

M

5.0

40

265

501

9

45

952

2.5

11

if. glabra

M

10.0

36

421

1309

17

50

1391

3.3

12

if. glabra

M

4.5

30

90

253

5

27

439

2.2

Clone
Number

*0ven-dry weight, excluding weight of fruit if present

Number
of
Sprouts

Average
Height
(feet)

63
operations should be an effective method of increasing sumac browse in
areas where most plants exceed six feet in height.

Summary
It was found that sumac is rather easily grown from seed, and that
it may be transplanted with a high rate of success.

Mowing was also

demonstrated to be an acceptable method of reclaiming, as deer browse,
clones which had exceeded the reach of deer.

The major findings of

this section are outlined below:
(1)

There was considerable interspecific variation in the size

and weight of sumac seeds:

about 46,000 seeds per pound of smooth

sumac; about 60,000 seeds per pound of staghorn sumac; and

about

52,000 seeds per pound of thehybrid sumac.
(2)

Scarification in concentrated sulfuric acid was required to

break seed dormancy, and the optimum length of time of scarification
for the hybrid sumac was 180 minutes.
(3)

Sumac was grown successfully from scarified seeds completely

covered with soil and planted at depths of one inch or less.
(4)

Seedlings grown under greenhouse conditions in soil trays and

Jiffy-7 peat pots were successfully transplanted to a nursery site at
the Tree Research Center.

The peat pot plants were transplanted intact

while those from the soil trays were planted bare rooted.

Two months

after transplanting, those transplants from the soil trays had a slightly
higher rate of survival although those in the peat pots had a signifi­
cantly greater degree of stem and root development.
(5)

Two-year-old sumac plants were transplanted from one site to

another several feet away.

Three transplant methods were employed:

planted bare rooted with minimal root exposure to air; planted bare

64
rooted after exposing the roots to air for one hour; and planted balledin-so±l with minimal disturbance to the roots.

All transplant methods

were successful, with the balled-in-soil method being the most successful.
(6)

Sections of sumac clones in which the height of many plants

exceeded six feet were cut at a height of four inches from the ground.
All clones resprouted vigorously, and it was suggested that this method
be used to lower the browse zone of tall sumac clones.

CONCLUSIONS

The results of this study indicate that sumac is a nutritious and
heavily utilized deer browse in northern Lower Michigan.

The importance

of sumac on the study area is underscored by the fact that it is both
highly productive and abundant in forest openings.

The plant is easily

cultivated in the nursery, and should respond well to establishment
under favorable field conditions.
Because of the importance of sumac as a deer browse, it is sug­
gested that planting sumac be considered as a part of a balanced vege­
tative management plan in the following situations on State and
Federally owned forests in Michigan:

1.

Along the berms of forest roads,

2.

In clearings and old fields designated as wildlife openings,

3.

In openings along the peripheries of established pine plan­
tations , and

A.

In newly planted and recently clearcut conifer plantations.

The last suggestion would be especially desirable as these areas
are often nearly devoid of deer browse.

It seems probable that such

treatments could be Integrated easily into a multiple-use forest manage­
ment concept, and in many cases they would add appreciably to the
wildlife value of commercial timber operations.

65

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APPENDIX

APPENDIX
INDEX TO COMMON AND TECHNICAL NOMENCLATURE

Vertebrate Animals

B.

C.

Technical Name:

Common Name:

Bonasa umbeZZus
CoZlnus vlrglnlanus
Lophortyx c a Z l f o m l o a
MeZeagrls gaZlopavo
Odocol Zeus vlrglnlanus
Pedlooetes phaslaneZZus
Phaslanus coZchlaus
SyZvlZagus fZorldanus
S a l u m s nlger
Tyrupanuahus aupldo

Ruffed grouse
Bobwhite quail
California quail
Wild turkey
White-tailed deer
Sharp-tailed grouse
Ringed-neck pheasant
Cottontail rabbit
Fox squirrel
Prairie chicken

Arthropod Animals
Technical Name:

Common Name:

Erlophyes rhols
HoZooaera chaZaofronteZZa
Idlomaoromerus blmiauZupemli
MeZaphls rhols

...
...
...
...

a
a
a
a

mite
lepidopteran
chalcid fly
louse

Woody Plants
Technical Name:

Common Name:

Acer rubrum
Acer saeeharum
AmeZocnchler arborea
Fragarla vlrginlana
Fraxlnuo amerlcana
Fraxlnus nigra
Plcea gZauaa
Plnus bankslana
Plnus reslnosa
Plnus strobus
PopuZus grandldentata
PopuZus tremuZoldes
P m n u s serotlna
P m n u s vlrginlana

Red maple
Sugar maple
Downy serviceberry
Virginia strawberry
American ash
Black ash
White spruce
Jack pine
Red pine
White pine
Big-toothed aspen
Quaking aspen
Black cherry
Choke cherry
73

74
Red oak
Dwarf sumac
Smooth sumac
Staghorn sumac
Common blackberry
Upland willow
Northern white cedar
American basswood
Poison ivy
Eastern hemlock
American elm

Quercus rubra
Rhus copallina
Rhus glabra
Rhus typhina
Rubus allegheniensis
Salix humilis
Thuja occidentalis
Ti.Ua amerioana
Toxicodendron radicans
Tsuga canadensis
Ulmus amerioana
D.

E.

Herbaceous Plants
Technical Name:

Common Name:

Achillea millefolium
Ambrosia artemisiifolia
Anaphalis margaritaoea
Anemone cylindrica
Antennaria plantaginifolia
Apocynum cannabium
Asclepias syriaca
Centaurea maculosa
Erigeron annuus
Erigeron canadensis
Hieracium aurantiacum
Hieracium florentium
Hieracium gronovii
Hypemicum punctatum
Physalis heterophylla
Prunella vulgaris
Ranunculus septentrionalis
Rumex acetocella
Solidago canadensis
Solidago sp.
Tragopogon dubius
V i d a cracca

Common yarrow
Common ragweed
Pearly everlasting
Thimbleweed
Plantain-leaved everlasting
Indian hemp
Downy milkweed
Spotted star-thistle
Wandering fleabane
Horseweed
Devil’s paint brush
Florentine hawkweed
Hawkweed
Common St. Johns-wort
Downy ground-cherry
Common self-heal
Swamp buttercup
Sheep sorrel
Canadian goldenrod
Goldenrod
Go a t ’s beard
Tufted vetch

Ferns, Mosses, Lichens, Fungi

Rusts

Technical Name:

Common Name:

Ceratodon purpureus
Cladonia arbuscula
Cladonia chlorophaea
Cladonia cristatella
Cladonia pyxidata
Cryptodiaporthe sp.
Fusarium sp.

...
...
...
...
...
...
...

a
a
a
a
a
a
a

moss
lichen
lichen
lichen
lichen
fungus
fungus

75

PhysaZospora sp,
PiZioZaria sp.
PoZytriaum juniperZnum
Pteridium aquiZinum
Pythium sp.
Sphaerotheca sp.
VertieiZZiwn sp.

. .. a fungus
... a fungus
,., a moss
.firacken fern
,.. a fungus
... a fungus
. .. a fungus

VITA
Hanley K. Smith
Candidate for the Degree of Doctor of Philosophy

Final Examination:

September 24, 1970

Guidance Co m m i t t e e : D r s . L . W . Gysel (Chairman), G . Sch n e i d e r , D . E .
Ullrey, and D. P. White

Dissertation:
The Biology, Wildlife Use and Management of Sumac In
the Lower Peninsula of Michigan
Biographical I t e m s :

Born October 28, 1940, Washington, D.C.
Married Eleanor L. Haag, March 23, 1968

Education:
Tulane University, BS, 1963
Texas ASM University, MS, 1966

Michigan State University, Ph.D., 1970
Experience:
Zoo Keeper, New Orleans Zoo, New Orleans, Louisiana, 1962
Research Assistant, Texas A&M University, 1964-1965
Research Assistant, Michigan State University, 1966-1969
Ecologist, U.S. Army Engineer District, St. Louis, Missouri
Organizations:
American Fisheries Society
American Society of Mammalogists
Wildlife Society