UNIVERSITY OF CALIFORNIA

COLLEGE OF AGRICULTURE

AGRICULTURAL EXPERIMENT STATION

BERKELEY, CALIFORNIA

Composition of

Common California Foothill Plants as a

Factor in Range Management

AARON GORDON and ARTHUR W. SAMPSON

Foothill Kange of the San Joaquin Valley

BULLETIN 627

March, 1939

UNIVERSITY OF CALIFORNIA
BERKELEY, CALIFORNIA

CONTENTS

PAGE

Review of literature 4

The San Joaquin Experimental Range 8

Sampling of species and analytical methods 12

Composition of graminaceous species 16

Organic constituents 17

Inorganic constituents 24

The calcium-phosphorus ratio 35

Evaluation of the species as forage 35

Composition of grasslike species 37

Organic constituents 39

Inorganic constituents 39

Evaluation of the species as forage 40

Composition of the broad-leaved herbs 40

Organic constituents 40

Inorganic constituents 54

The calcium-phosphorus ratio 61

Evaluation of the species 61

Composition of browse from shrubs and trees 64

Half-shrubs 64

Nondeciduous species 72

The calcium-phosphorus ratio 73

Evaluation of the species 74

Comparative levels of constituents in the six plant groups 75

Application to range management 81

Comparative forage values of the different growth stages 82

Role of weeds in a range community 84

Soils in relation to plant types and forage utilization 85

Criteria for judging the value of a forage plant 87

Shortening of growing season as affecting nutritive values 87

Chemical values and grazing capacity 88

Summary 89

Acknowledgments 91

Literature cited 92

COMPOSITION OF COMMON CALIFORNIA

FOOTHILL PLANTS AS A FACTOR IN

RANGE MANAGEMENT1 2

AARON GORDON3 and ARTHUR W. SAMPSON*

This bulletin reports a study of chemical characteristics of certain foot-
hill range plants commonly found in the interior valleys and on the coastal
strip of California. Information obtained from such investigations should
point the way to a more scientific approach to range revegetation and
maintenance, and to a more accurate evaluation of the carrying capacity
of the range.

In half a century of grazing by domestic livestock conspicuous changes
have taken place in the plant population of this vast grass-woodland
area. In many instances the carrying capacity has declined sharply, and
a portion of the top soil has been removed. Evidence points to the fact
that a substantial proportion of the original perennial graminaceous
species has been replaced by annual exotic grasses and other herbs of
southern Europe ; this accounts for the inclusion of many such species
in this study. So conspicuous has been this transformation that future
grazing management plans must be focused largely upon this relatively
early-maturing vegetation.

Although various grazing systems have been employed on some of
these ranges with a view to improving the forage crop, uniformly satis-
factory results in revegetation have not been obtained. Better under-
standing of the plant-nutrition problem is obviously basic to the
improvement and maintenance of the forage cover. To understand the
continuous transformations in the range-forage complex one must rec-
ognize the changes peculiar to the life cycle of individual species and the
departures from these changes caused by the various limiting factors,
including different degrees and character of pasture utilization.

The study was undertaken in 1935, on the San Joaquin Experimental
Range of the United States Forest Service, a branch station of the Cali-
fornia Forest and Range Experiment Station located near O'Neals.

1 Received for publication June 19, 1938.

2 Results of a cooperative investigation conducted by the California Forest and
Range Experiment Station of the United States Forest Service, United States De-
partment of Agriculture, and the California Agricultural Experiment Station.

Acknowledgment is also made to the Works Progress Administration for assistance
rendered through its Official Project No. 465-03-3-630, Unit B-12.

3 Associate Plant Ecologist, California Forest and Range Experiment Station,
United States Forest Service.

4 Associate Professor of Forestry and Plant Ecologist in the Experiment Station.

[3]

4 University of California — Experiment Station

The results are reported under the following general headings: (1)
composition of the chief grass species composing the different subtypes ;
(2) composition of the grasslike species; (3) composition of the broad-
leaved herbs, shrubs, and arborescents ; (4) chemical characteristics and
phylogenetic relations; (5) habitat and seasonal influences on composi-
tion ; and (6) application of the results to range management.

REVIEW OF LITERATURE

It is common knowledge that the characteristic chemical composition of
a species is measurably influenced by the habitat. Some investigators
have also reported that the chemical levels of pasture plants vary from
the earliest inception of growth to maturity.

It is fairly well established that the acidity of a soil is an important
factor in rendering nutrients available to vegetation. Davis, Hoagland,
and Lipman (9)5 concluded that absorption of ions is influenced by the
amount of carbon dioxide liberated by the roots. Pierre and Pohlman
(39), using corn, sorghum, and sudan grass, found that differences in
soil acidity from pH values of 4.60 to 6.60 do not affect the concentration
of calcium and phosphorus in the sap of these three species. Kauter (28)
found that the percentages of phosphorus, calcium, and potassium in the
ash of hay increases with higher pH soil values. This he states is due to
dominance of species high in these constituents in soils of lower hydrogen
ion concentration. Hoagland (24), working with barley in water cul-
tures, reported that the values of phosphorus, and less conspicuously of
calcium, decrease with higher pH. Truninger and Grunigen (48) found
no relation between the soil reaction in pll ranging between 4.5 and 8.0
and the content of phosphoric acid, potassium, and calcium in forage.

The addition of soil fertilizers generally appears to affect plant com-
position. Hissink (23) concluded that while the proportions of mineral
constituents in the plant depend primarily on the species concerned, they
are also affected by the minerals present in the soil. Cooper (5) reported
that the ash constituents of pasture grasses studied are largely deter-
mined by the amounts of the various constituents available in the soil.
He found that plant nitrogen and ash decrease as the soils become de-
pleted. Jacob (27) found that addition of potassium to the soil increases
the potassium content of the forage in many cases, but reduces the nitro-
gen and lime contents. Mortimer and Ahlgren (36) , in studies with Ken-
tucky bluegrass, reported that fertilization with nitrogen decreases the
calcium and phosphorus content of this species, whereas addition to

5 Italic numbers in parentheses refer to "Literature Cited" at the end of this bul-
letin.

Bul. 627] Composition of Foothill Plants 5

the soil of phosphorus and potassium increases the nitrogen content of
the herbage. They also stated that phosphate fertilization increases this
constituent of the grass. Truninger and Grunigen (48) found a reduc-
tion of lime content in the plant when excessive amounts of potassium
were added to the soil. Daniel and Harper (<§), in studying the composi-
tion of Andropogon furcatus, concluded that the study of a single nutri-
ent element in the soil will not give a reliable indication of the amount of
that element found in the plant.

That drought influences the composition of forage has been pointed
out by various workers. Ferguson (12) states that in forage there is a
decrease of calcium and phosphorus during a period of drought. "Wood-
man, Norman, and French (52), as well as others (29), found that the
most conspicuous effects of drought are : a decided decline in the per-
centage of protein ; a slight increase in the percentage of nitrogen-free
extractives and crude fiber ; an abrupt rise in the percentage of calcium,
accompanied by a decline in the percentage of phosphorus ; and a pro-
nounced reduction in moisture content with a premature onset of ligni-
fication. Franklin (14) reported a fairly close relation between variation
in the mineral content of pasture forage and certain meteorological data.
Watkins (50) found that, in range grasses of the Southwest, phosphorus
is more commonly deficient than calcium.

In various stages of plant development, movement of elements takes
place from one plant organ to another, and from the roots to the soil.
Thus Burd (3) reported an orderly increase in nitrogen and potassium
in barley during rapid growth, followed by losses at the time of heading.
Murneek (37) found that tomato plants absorb the greatest quantities
of nutrients and synthesize the largest amounts of organic substances
when fertilization of the flowers was permitted, but when the fruit was
not allowed to develop. Murneek (38) later stated that decline in vege-
tative growth, which is associated with the reproductive period, is ac-
counted for by the withdrawal of nitrogenous products from the leaves.
Austin (1, 2), studying the soybean, found that carbohydrates accumu-
late in deflorated plants, but that the intake of hydrogen ion and minerals
is simultaneously curtailed. Richardson, Trumble, and Shapter (40),
working under controlled conditions with the perennial, Phalaris tuber -
osa, reported that, at plant maturity, nitrogen and phosphorus descend
to the basal portion of the stem and to the roots, but a substantial portion
of the total potash of the plant may diffuse from the roots into the soil.

The different parts of the plant may vary in composition. In the study
of forest trees Seiden (45) reported that the ash content is lowest in the
leaves of the older branches nearer the ground. In a more comprehensive

6 University of California — Experiment Station

study McHargue and Roy (34), working with 23 species of deciduous
trees, found that the young foliar organs have the highest percentage of
phosphorus, potassium, and nitrogen, whereas the mature foliage con-
tains the highest percentage of ash, silicon, and calcium. Sampson and
Samisch (44) , working with Quercus Gambelii and Q. Kelloggii, reported
high content of crude protein and crude fiber in the young leaves, but as
the foliage expands the amount of these constituents per unit area de-
creases. Stapledon, Fagan, Evans, and Milton (47) reported that in
Italian rye grass, the young leaves, and especially the leaf sheaths, are
nearly twice as rich in protein as the stem. Silica-free ash is also higher
in the leaves.

Conspicuous changes in plant composition take place as the season
advances. Various investigators (10, 25, 31, 40) have found that with ad-
vancing maturity of forage plants the crude protein content decreases
and the crude fiber and nitrogen-free extracts increase. Hart, Guilbert,
and Goss (22) found that the dry matter in the forage varies from high
protein content in early growth stages to that of poor roughage when
the herbage has dried. Bur clover, however, retains a relatively high
nutritive value throughout the year, whereas the nutritiveness of most
annual species is short-lived. McCreary (33) reported the highest nitro-
gen content in ungrazed pasture grass during the succulent stage, fol-
lowed by decline to maturity. Sampson and McCarty (42) , working with
Stipa pulchra, found definite seasonal trends in the annual march of
carbohydrates, with a decline during the period of rapid growth and a
rise after the season's peak of development. Roberts (41) found that the
ash content of some native perennial grasses studied increases with ma-
turity, whereas in some exotic grasses the ash content decreases later in
the season. Several other investigators (3, 40, 51) have reported high
relative amounts of certain elements, notably potassium, phosphorus,
and nitrogen, during the early growth stages, followed by decrease as
maturity approaches. In the study of Phalaris tuberosa, Richardson,
Trumble, and Shapter (40) found the crude protein to be 33 per cent in
the tillering stage, with a decline to 3.37 per cent at maturity. The levels
of phosphate and potash content decrease, while the percentage of crude
fiber increases continuously from the early leaf stage to maturity. Green-
hill and Page (17) reported appreciable fluctuation in calcium content
in grasses during the growing season, whereas phosphorus had a definite
seasonal trend, falling during drought and the early summer "flush"
period, and recovering after each decline. Van Itallie (49), who worked
on the chemical composition of species of grasses at various stages of
growth, reported that the nitrogen, fat, and mineral constituents,

Bul. 627] Composition of Foothill Plants 7

with the exception of silicic acid, decrease with advancement of the
season, whereas the crude fiber content increases steadily. Fagan and
"Watkins (11) found that herbs and grasses decline in nutritive values
with advancing maturity. Sampson and Parker (43) noted that in St.
Johnswort (Hypericum perforatum) there is a rapid rise in sucrose until
blossoming time, after which the amount falls off steadily.

The levels of composition during growth vary with the species. Mc-
Creary (33) found that plants of high altitudes contain more nitrogen
than those growing at lower elevations, when collected in the same growth
stage. Roberts (41) reported that western forage plants increase in feed-
ing value with increasing altitude. Van Itallie (49) noted that although
the trend of the constituents is similar in all grass species studied, the
relative amounts in the different species vary. Thus Holcus lanatus and
Alopecurus pratensis were high in ash and potassium content, while Fes-
tuca rubra and Poa pratensis were low. Du Toit, Louw, and Malan (10)
found pure species, grown in the same soil and exposed to the same cli-
matic conditions, show appreciable differences in mineral and protein
contents when harvested at definite intervals. Truninger and Gruni-
gen (48) state that the mineral content of meadow forage, within certain
limits, is influenced more strongly by the botanical composition than by
soil fertilization. Pure grass stands show a smaller content of phosphoric
acid and lime than do clover-grass meadows. Forbes, Whittier, and Colli-
son (13) concluded that seasonal variations in ash content depend largely
upon habitat rather than on the plant species. Daniel (7) contends that
the particular plant species is much more important than either the soil
or the land treatment in determining the mineral content. He found that
legumes contain more calcium than grasses. In legumes there is a high
calcium-phosphorus ratio, whereas in grasses this ratio is relatively low.
Kauter (28) concurred with these results. Richardson and associates
(40) in their investigation of the mineral content of pastures, concluded
that superphosphate applied to natural pasture induces changes in bo-
tanical composition of the pasture and in plant succession. McHargue,
Roy, and Pelphrey (35) , found appreciable variations between species in
the amounts of iron, copper, zinc, or manganese. Capen and LeClerc (4) ,
working on Alaska species, found measurable differences in the ash and
sugar content in forage species.

Leaching of certain components may affect the composition of the
plant. Bur clover, according to Guilbert and Mead (18), is rich in pro-
tein, and has a narrow nutritive ratio. This ratio in mature vegetation is
affected by rain, which these authors report causes the soluble constitu-
ents to be leached out. Guilbert, Mead, and Jackson (19) found that the

8 University of California — Experiment Station

ingredients most susceptible to leaching from the plant are those of cer-
tain constituents of the silica-free ash, and of the nitrogen-free extract.

The degree and season of pasturing have also been shown to affect the
composition of the forage. Hopper and Nesbitt (25) reported that fre-
quent clipping tends to maintain succulence in forage, a condition associ-
ated with high crude protein and low crude fiber. Harrison (21) found
that removal of top growth lessened the carbohydrate storage, and if the
clipping were sufficiently severe, no carbohydrates were stored. Mc-
Carty (32) obtained similar results in his studies of Avena fatua. Graber
and associates (15, 16) found that the storage organs of a plant clipped
at maturity contain more organic material than those of a plant har-
vested during the growth period.

To summarize : Analytical studies of the various constituents, particu-
larly in regard to inorganic elements, reveal that much confusion and
speculation exist. The inconsistencies, even as to general trends in the
mineral constituents, and often in the organic contents, leave few defi-
nite conclusions. That there is high protein content and low fiber and
nitrogen-free extract in very early growth stages, followed by gradual
decline in the former and increase in the latter two, with the advance
of the season, has been fairly well established. The quantitative trend in
the ash content, and in the essential constituents thereof, has been little
clarified. Moreover, there remains to be determined the effect of herbage
removal on the physiological reaction of the plant, as measured in chemi-
cal constituents.

The divergent results reported in the literature are ample justification
for undertaking the present study.

THE SAN JOAQUIN EXPERIMENTAL RANGE

The San Joaquin Experimental Range of some 4,700 acres, where the
plant collections were made, appears to be fairly representative as to
soils, vegetation, and climate of the foothill area of the region generally.
The soils" of the area are nearly all residual in character, being derived
from decomposition and disintegration of coarse-grained granodiorite
rock. They are of sandy loam or coarse sandy loam texture. Many stones
are present as outcrop and as loose stones throughout the soil profile. Bed-
rock occurs at a depth of a few inches to 3 feet from the surface, with an
average depth of about 18 inches. These soils are low in organic matter
and close to neutral in reaction. Where undisturbed, there is usually a
darkened organic surface layer about 1 inch in thickness, in which abun-

6 Eeported through the courtesy of Dr. E. E. Storie of the Division of Soil Tech-
nology, University of California.

Bul. 627]

Composition of Foothill Plants

9

dant grass roots are located. There is little evidence of a true subsoil. The
soil material above the bedrock is of about the same texture as that of
the surface but contains less organic matter and is of lighter color. Thus
the residual soils classify as Visalia sandy loam, rock outcrop phase.
Small alluvial areas, 100 to 200 feet in width, occur in narrow valleys.
These soils are of dark brownish-gray color and also correlate with the
Visalia sandy loam or Visalia loam types.

The vegetation is essentially an open oak savanna, broken here and
there by clumps of brush and small arborescents, of which Ceanothus,

Fig. 1. — In the foreground is seen a swale which supports a dense cover of
grasslike plants. Such areas contribute richly to the forage supply after the
adjoining grass cover has dried.

Aesculus, Sambucus, Rhamnus, silverleaf (Lupinus) , and deerweed
(Lotus) are conspicuous (frontispiece and fig. 1). Quercus Douglasii is
the most characteristic tree, with Q. Wislizenii and Pinus Sabiniana
conspicuous in restricted sites. The herbaceous cover is composed largely
of annual grasses, with an abundance of broad-leaved annual herbs.7

The woody plants named occur with the grass dominants throughout
the range, and in an occasional restricted area they form the major vege-
tation. The several swales support a dense cover (60-90 per cent density)
mainly of Eleocharis palustris, Juncus bufonius, and J. oxymeris (fig.

7 Unpublished studies by the California Forest and Eange Experiment Station re-
veal the fact that on 2,165 sample plots scattered at random throughout the San
Joaquin Valley ranges, annuals now account for 97 per cent of the herbage, with
bunch grasses and other perennials accounting for only 3 per cent.

10

University of California — Experiment Station

1). Much of this vegetation remains succulent long after the grasses of
the well-drained soils have matured, when it furnishes important forage.
The various species analyses of which are reported in this bulletin are
here listed to indicate further the character of the cover :

Grasses and Grasslike Species8

Avena barbata Brot. (Slender wild

oats)
Bromus arenarius Labill. (Australian

chess)
Bromus mollis L. (Soft chess)
Bromus rigidus Roth. (Ripgut grass)
Bromus rubens L. (Foxtail chess)
Festuca megalura Nutt. (Foxtail

fescue)

Melica imperfecta* Trin. (California
melie)

Boa scabrella* (Thurb.) Benth. (Pine
bluegrass)

Eleocharis palustris* R. and S. (Wire
grass)

J uncus bufonius L. (Toad rush)

J uncus oxymeris* Engelm. (Slender-
leaved rush)

Broad-leaved Herbs8

Amsinckia Douglasiana DC. (Buck-
thorn weed)

Cryptantha flaccida (Dougl.) Greene
(White forget-me-not)

Er odium botrys Bertol. (Storksbill)

Erodium cicutarium L'Her. (Red-stem
filaree)

Erodium moschatum L'Her. (White-
stem filaree)

Filago gallica L. (Woolly filago)

Gilia tricolor Benth. (Birdseye gilia)

Godctia amoena (Lehm.) Lilja. (Sum-
mer's darling)

Godetia quadrivulnera (Dougl.) Spach.
(Spotted godetia)

Hemitonia virgata Gray (Yellow tar-
weed)

Hemitonia Wrightii Gray (Tarweed)

Layia pentachaeta Gray (Tidytips)

Lessingia germanorum Cham. (Lessin-
gia)

Linanthus ciliatus (Benth.) Greene
(Pink linanthus)

Shrubs

Aesculus calif ornica (Spach.) Nutt.
(California buckeye)

Ceanothus cuneatus (Hook.) Nutt.
(Wedgeleaf ceanothus)

Ceanothus divaricatus Nutt. (White-
bark soapbloom)

Lupinus albifrons Benth. (Silverleaf
lupine)

Lotus americanus (Nutt.) Bisch.

(Spanish clover)
Lotus strigosus (Nutt.) Greene (Hairy

lotus)
Lotus subpinnatus Lag. (Fine-leaved

lotus)
Lupinus Benthami Hel. (Bentham

lupine)
Lupinus bicolor Lindl. (Late valley

lupine)
Lupinus formosus* Greene (Wand

lupine)
Navarretia viscidula Benth. (Viscid

navarretia)
Plagiobothrys nothofulvus Gray (Pop-
corn flower)
Plagiobothrys tenellus Gray (Popcorn

flower)
Silene gallica L. (Windmill pink)
Tri folium tridentatum Lindl. (Tomcat

clover)

and Trees

Lotus scoparius (Nutt.) Ottley (Deer-
weed)
Quercus Douglasii H. & A. (Blue oak)
Rhamnus calif ornica Esch. (California

buckthorn)
Rhus diversiloba T. & G. (Poison oak)
Sambucus glauca Nutt. (Blue elder-
berry)

The climatic characteristics of the foothill range are typified by dis-
tinctly seasonal rainfall, by temperatures sufficiently mild to favor
growth of "winter" annuals, and by long, hot, dry summers.

Rains of sufficient magnitude to initiate seed germination and the

8 Species of grasses and broad-leaved herbs marked with an asterisk are of peren-
nial growth.

Btjl. 627]

Composition of Foothill Plants

11

greening of bunch grasses usually come in October. A rain of about 1
inch starts vigorous plant activity. Culmination of the rainy season is
generally about the last of April, but light showers are sometimes re-
ceived as late as July. Cumulative rainfall records compiled by the Cali-
fornia Forest and Range Experiment Station for the period of study
(fig. 2) show the annual precipitation to vary from 22.65 inches, as in
1935-36, to 29.60 inches, in 1934-35. In 1935-36 a dry period occurred

1 1 1 1 1 1 1 1 1 1

_ 29.60 TOTAL

1 1

-

p-J^ -22.91 TOTAL 22.65 TOTAL

-

-

J7f

i — V-T

-

J f t

-

LEGEND

1934-35

1935-36

-

-

1936-37

-

-

H —J1
. — irj-

-

._„/"

! i— '[r — '

K ,'. l i, '' rt-fii ,'.. '

' i ' * ' 1 ' ' ' ' ' 1 ' ' ' ' ' 1

i i i i i I i i i i i 1 . i . i . 1 . . i i i 1 i . i i i 1 i i i

^4-J

Fig. 2. — Cumulative inches of rainfall recorded on the San Joaquin Experi-
mental Range for the seasons of 1934-35, 1935-36, and 1936-37.

after favorable early October rains. This dry period was not broken in
December, hence the growth of early winter forage was below normal. In
1934-35, distribution of rainfall was highly favorable to growth from
its inception to its termination. Rains that fall early in May and there-
after, promote little or no growth and may actually decrease the nutritive
value of such forage as has matured. Variation in the total annual rain-
fall, and in its seasonal distribution, may be sufficient to influence meas-
urably from year to year the reproductive capacity and the yield of
forage.

The temperature factor, with an extreme annual range of approxi-
mately 100° Fahrenheit, is limiting to growth only from about December
to February (fig. 3) . During this period the mean minimum temperature
is little above 40°, with occasional periods of freezing temperatures. In
January, 1937, some of the herbage was frozen to the ground, whereas
in the two previous years the rate of growth was almost nil during the
several weeks of cold weather. In February, with the characteristic an-
nual rise in temperature the growth rate increases sharply. Maximum

12

University of California — Experiment Station

temperatures are reached in July and August, 2 or 3 months after the
annual grasses have matured. As seen from the graphs of the mean tem-
peratures for 1935, 1936, and 1937, there is an orderly rise and fall,

100

1 1 1 | 1

"

1 1 | 1 1 1 1 1

1 1 1 1 M i

i

i i | i i ii i | i I i ii | i i i i i | ii I i i | r i

1 1 1

90

k

/\ >■

-

80

-

/ \/V"/\

&w

-

2

X 70

2
UI

a.

X

560

w
w

a

O 50

u

Q

-

A '

/V

' 1/ v Y\ /\

v y wl \

-

"

/\ ik.*-

wA f\

\ i/' N

JrJ

\ /V' v"

A"

40

-"\

i v

MEAN AT TWO-HOUR INTERVALS
1935

Nv\

-

1937

-

MllllMIMlllllll 1 M 1 1 1 1 1 1 1 1 1 1 1 1 ! 1 1 1 1 1 1 1 1 1 1 1

Fig. 3.

-Temperatures recorded on the San Joaquin Experimental Range
for 1935, 1936, and 1937.

respectively, in temperatures before and after the summer maxima. Be-
tween June and October the succulent vegetation consists of browse,
species of Lotus, and small patches of grasslike vegetation. Departures
in mean temperatures are seen to be insignificant for the past two years.
If snow falls at all, it usually melts soon.

SAMPLING OF SPECIES AND ANALYTICAL METHODS

Studies of range plants are complicated by the fact that the species which
compose the cover are not in a uniform developmental stage at a given
sampling period. In any habitat some species germinate or vegetate
early ; they may have virtually completed the growth cycle when later-
maturing species start growing vigorously. Individuals of the same
species often occur in different growth stages at a given period. Accord-
ingly, random sampling without the recognition of the stage of develop-
ment, or the collection of composite samples, has proved of little value
in building up information leading to better understanding of range
management. It is only through the accumulation of knowledge of the
composition of individual species, taken at definite physiological stages,
irrespective of chronological periods, that reliable data may be obtained.
The species studied were taken on plots characteristic of different soil

Bul. 627]

Composition op Foothill Plants

13

and drainage conditions. Samples of most species were collected on a
small plateau of about 6 acres (fig. 4). In addition, several species were
grown from seed in the plant nursery of the range, which is a test plot
(fig. 5) of forage species, the soil of which is uniformly deep and well

Fig. 4. — Portion of the main range area where about 70 per cent of the species
studied were collected in various growth stages. The herbage, with an estimated
density of 35 per cent, consists mainly of Festuca megalura, Erodium ootrys,
Bromus rigidus, B. mollis, and Avena haroata. The arborescents seen are Aesculus
calif ornica, Quercus Douglasii, and Q. Wislisenii.

drained. Also a few species were collected at the ranch headquarters on
undisturbed soil.

Detailed record was made of the particular growth stage of each species
at the time of sampling. Collections were made according to stages of
plant development rather than chronological date. The composite sam-
ples for the analyses were procured from several plants collected at ran-
dom and all of as nearly the same growth stage as possible. Effort was
made to secure the material in the sequence here listed :

1. Early leaf stage (initial aerial growth) .

2. Just before flowering.

3. Plants in full bloom.

4. When the seeds were in dough stage (endosperm plastic) .

5. Plants mature, seeds cast.

6. Herbage dry and weathered.

These growth stages are presented in figures 6 and 7.

14

University of California — Experiment Station

Occasionally in the same habitat, and on the same date, two growth
stages of some of the species were procured. The samples of all herbace-
ous species were cut with shears about 1 inch from the ground surface.

Fig. 5. — Areas 6 x 20 feet were seeded in the nursery to facilitate procur-
ing samples of very young herbage and also of the more advanced growth.
A, Bromus mollis; and B, Plagiobothrys tenellus.

The samples of all species of the rosette form, and the earliest collections
of grasses and grasslike plants, were washed in several changes of water
to remove surficial soil particles and other foreign materials. Prior to the
adoption of this procedure, analyses were made of washed and of care-
fully wiped but unwashed samples, and these analyses revealed that
washing did not influence the chemical composition of the succulent ma-

Bul. 627]

Composition of Foothill Plants

15

terial. Samples of trees and shrubs consisted chiefly of the foliage, which
was stripped off by hand ; but in several species flowers, fruit, and cur-
rent branch growth were collected and analyzed separately. At the time
of collecting, additional specimens were placed in a plant press for sub-
sequent observation and verification of the particular growth stage which

Fig. 6. — Avena barbata showing five of the growth stages from which the sam-
ples were collected. A, Early leaf stage; B, just before flowering; C, full bloom;
D, seeds in dough stage ; E, herbage weathered and the seeds cast.

each sample represented; these specimens served to classify the data with
respect to march in composition. Several of the pressed specimens were
sketched to scale for permanent record.

The fresh weights of the samples were recorded immediately following
the collection, after which the material was dried in an electric oven, with
circulating air, at a temperature of 70° C. When thoroughly dry, the
samples were finely ground, bottled, and tightly stoppered. After the
analyses of the samples were completed an aliquot part of each sample
was dried at 100° C to constant weight ; the analytical data were then ex-
pressed on this moisture-free basis.

16

University of California — Experiment Station

In the determination for nitrogen, calcium, ash, and moisture, Official
and Tentative Methods of Analysis of the Association of Agricultural
Chemists" was followed. Phosphorus was determined colorimetrically, as
described by Kuttner and Cohen (30). The only modification made was
that the sample was ignited with MgO, dissolved in strong HC1, and after

Fig. 7. — Quercus Douglasii showing the growth stage when the samples "wore

collected. A, Early leaf stage; B, full bloom ; C, fruit mature but none cast.

rinsing the solution into a 100-cc volumetric flask, it was brought just to
neutral, as indicated by litmus, before the sulfuric acid and other re-
agents were added. Crude fiber was determined by the method described
by Sharrer and Kurschner (46). Silica was determined by treating the
ignited residue of the sample with concentrated HC1 and 60 per cent
HC104 and igniting the residue after digesting on the steam bath, and
then filtering.

COMPOSITION OP GRAMINACEOUS SPECIES

Table 1 gives the location where the samples of graminaceous species
were collected, the date of sampling, the stages of growth, and the results
of analyses. The first column, showing the respective growth stages, is
arranged in sequences from the early leaf stage to the time of full ma-

0 Association of Official Agricultural Chemists. Official and tentative methods of
analysis of the Association of Agricultural Chemists. Association of Official Agricul-
tural Chemists. Washington, D. C. 3d ed. 593 p. 1930.

Bul. 627] Composition of Foothill Plants 17

turity, for the respective years and locations. Where two samples repre-
senting a specific growth stage are given, the slightly younger sample is
presented first. Although definite growth stages are given for each sam-
ple, the fact must not be overlooked that growth is a continuous process,
hence the stages designated are necessarily somewhat approximate.

There is a continuous and rather orderly change in composition of all
constituents here reported, from the earliest appearance of leaf blades to
plant maturity. The content of most constituents, with the exception of
crude fiber, is highest in early growth and declines rather uniformly,
although at a different rate in the various stages, reaching the lowest
point in most species at full maturity.

For each of the three years of study there is an impressive consistency
in the composition of all the grass species for any given growth stage. It
is well known that the character of the soil medium may affect absorption,
and therefore the composition of the plant. However, under the range
conditions concerned, the soil solution is evidently in an approximate
state of equilibrium and does not appear measurably to affect the com-
position of the supporting vegetation from year to year.

Organic Constituents. — The uniformity in composition with respect
to organic constituents at any given growth stage is illustrated in figures
8 and 9, which give, for three years, the march in composition of the pro-
tein and fiber of the annual species, Avena barbata, and the perennial,
Poa scabrella. In both these constituents by far the most precipitous
change occurs from the early leaf period to the time that the plant is in
late bloom, designated as A B in figures 8 and 9. From the time of full
bloom to maturity, designated as B C in these figures, the rate of change
is slight and inconspicuous, though the trend is in the same general direc-
tion. When the plant first reaches maturity, until it is thoroughly weath-
ered in the late summer, designated as C D, there is little change in the
contents of crude protein and crude fiber. This fact is impressive in view
of the lapse of time between procuring samples of the recently matured
plant, indicated by C in the figures, and the older, bleached sample, desig-
nated by D. Obviously this behavior applies only to those species whose
seeds are not scattered between collections of the samples. In Bromus
mollis, whose seeds are mostly retained long after maturity, the relatively
large changes in protein content after the herbage has dried, are ac-
counted for by the proportion of the seeds retained. On June 13, 1936,
and on June 14, 1937, for example, when only a few of the seeds had been
cast in the field samples, the percentages of protein were 6.10 and 6.25,
respectively. On September 10, 1936, when most of the seeds had been
cast, the percentage of protein was 3.76.

18

University of California — Experiment Station

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24 University of California — Experiment Station

Where change in composition of various stages is discussed in this
report the composition of the earliest leaf stage is taken as unity, or 100
per cent. On this basis, calculations from table 1 bring out the fact that
the crude protein content in Bromus mollis and in Avena barbata, from
the earliest leaf stage to maturity, when most or all of the seeds are cast,
declines 83 per cent in 1936 and 89 per cent in 1937, respectively. In
Melica imperfecta, a perennial species, the decline is 87 per cent for the
corresponding growth stages. In contrast, the crude protein levels at the
early growth stage for the various grass species do not differ greatly.
Calculations show the difference between the highest and lowest protein
content for all grasses reported, when in the early leaf stage, to be 44 per
cent, whereas there is a difference of 94 per cent in A. barbata for the
entire growth cycle, as in 1935.

The crude fiber, as found by many workers, is lowest in the early
growth of the plant and highest at the time of full maturity. In Avena
barbata, in 1937, the crude fiber content increases 115.6 per cent from
the early leaf stage to full maturity ; the other species investigated show
similar increases for corresponding growth stages. In contrast to the
wide seasonal variation within a species there is small difference in this
constituent among the species studied for a given growth stage.

The variations in the level of crude fiber in Avena barbata at five
growth stages for three successive years are shown in figure 10. The crude
protein content is included in this figure for purposes of comparison. The
averaged composition of these two organic constituents at the early leaf
stage is shown in figure 11 for six grasses. Comparison of figures 10 and
11 shows that the differences in the percentages of crude fiber and crude
protein between the various species investigated are not so great in any
one growth stage as those in a single species for the entire season of
growth.

Inorganic Constituents. — "Without exception the mineral content of
all the grass species here reported is highest in the very young herbage.
Although the decline is not so precipitous as that of crude protein, the
mineral constituents decrease in most species with aging of the plant,
generally reaching the lowest point when the top growth has dried.

The march of the ash content is similar to that of the potassium in
Avena barbata and Poa scabrella, as shown in figures 12 and 13. The
highest levels of these components occur in the very young herbage, and,
like the protein content, they decline rather sharply to the late blossom-
ing period, as shown by the segments A B. From the latter period of ma-
turity, indicated by B C, these constituents are fairly constant. Moreover,
from maturity to thorough weathering — a dry, hot period from May to

Bul. 627]

Composition of Foothill Plants

25

35

30

25

i 1 1 r

B
p-' "".■•■•/" '

FIBER

16

20 h6 /

6
18

LEGEND

1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

20

15 -

10 -

-

1 1 1

III 1

-A

LEGEND

lk\

V

1Q3^ FTFI.D PTPT

\

1936 FIELD PLOT

*

1937 FIELD PLOT

-

>\ <?

-

-

\\\\

-

-

V V

-

-

V \
V \

PROTEIN

-

V \

V •• \
v \ \
v '• \

-

—

V ■ \

—

-

\\

B

'"°--..

-

15^

*i^>^:-.

-

^?5^-^ c d:

** ^^l^^T***^^ ° -

-

i i i

iii i

EARLY OUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY
FLOWERING STAGE DRY

PLANTS DRY

AND
WEATHERED

Fig. 8. — March of crude protein and crude fiber in Avena barbata, an annual
grass, for 1935, 1936, and 1937.

26

University of California — Experiment Station

40-

U

w

I 1

1 1

1 1

— 1 ■ .-6

D

-

/ c

© ■

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x .•■ i?

FIBER

-

LEGEND

1Q3*>

FIELD PLOT

1936

FIELD PLOT

1 1

1937

1 1

FIELD PLOT

6-1

w

O 15

10 -

1 1 1 1 1 1 1

LEGEND

193FJ Firm pt.dt

\ a 1936 FIELD PLOT

. \ 1937 FIELD PLOT

\
\

\

: \\ :

•. \
••. \

•A

"^V PROTEIN :

Y\B :

\^'*-

----^ C D ~

^oi-^a.---— 0 -

-© -

-

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 9. — March of crude protein and crude fiber in Poa scabrella,
a perennial grass, for 1935, 1936, and 1937.

Bul. 627]

Composition of Foothill Plants

27

September — the levels of ash and potassium, as shown in segment C D,
also remain practically constant. The greater smoothness of the curves
representing the nursery samples is accounted for by the even age of
the plants.

It is seen from table 1 that the highest silica-free ash content recorded
for any grass species is 12.92 per cent, and the lowest is 1.61 per cent. In

40

Avena barbata

1 . Early leaf stage

2. Just before flowering

3. Full Bloom

4. Seeds in dough stage

5. Plants mature.

seeds cast

15

FIBER PROTEIN

Fig. 10. — Crude protein and crude fiber in Avena harhata

for five growth stages for 1935, 1936, and 1937.

the field plot of Bromus mollis, in 1936, the decline in ash content from
the early leaf stage to maturity is 75.4 per cent ; in Festuca megalura it
is 74.7 per cent. The seasonal differences shown are representative of the
grasses studied.

When the same growth stage of the various species is compared, the
difference in the amount of ash is not conspicuous. When, for example,
the highest and the lowest silica-free ash contents of all the grass species
are compared for a given growth stage, the percentage differences are :
in the early leaf stage, 60.9 per cent ; at time of blossoming, 57.9 per cent ;
and at maturity, 70.0 per cent. The seeds of the grasses are very low in
silica-free ash, ranging from 2 to 3 per cent.

The potassium content shows a general downward trend throughout
the growing period, most of the species reaching the lowest values at full
maturity. In Avena barbata (fig. 12) the potassium content increases
slightly from the time the seeds are in the dough stage to maturity. This

28

University of California — Experiment Station

behavior is probably accounted for by the fact that a portion of the first
spring leafage matures early and is lost under field conditions, and is,
therefore, not available for inclusion in the sample. Calculations made
from table 1 give a maximum difference between the highest and lowest
content of potassium in A. barbata for the three years reported of 66.3
per cent, and in Bromus mollis of 80.1 per cent. In the other species
studied similar variations in potassium content are found.

25

1. Avena bartata

2. Bromus mollis

3. Bromus rigidus

4. Festuca megalura

5. Melica imperfecta

6. Poa scatrella

15

PROTEIN

FIBER

Fig. 11. — Crude protein and crude fiber in the early leaf stage
of some of the grass species studied.

The potassium levels for a given growth stage show little difference in
any of the species for the three years of study. Also, the levels of this
constituent are seen to differ less in any specific growth stage for all the
grass species investigated than for a given species for the entire growing
season. In the early leaf stage of Avena barbata in 1935, 1936, and in
1937, there is a variation in the potassium content of only 34.9 per cent,
and in Bromus mollis for 1936 and 1937 of 14.5 per cent. A similar rela-
tion holds for the other species studied.

The preceding discussion, based as it is upon three successive years of
study of eight dominant annual and perennial grass species, is convinc-
ing of the fact that analytical data of a species whose sample is not rep-
resentative of a specific growth stage, has little or no bearing on the

Bul. 627]

Composition of Foothill Plants

29

8-

1 1 1

ill 1
LEGEND

\A

lor^fi >jrrpc.pDy

1Q3R FTFTX) PT.OT

_ \

1936 FIELD PLOT

1937 FIELD PLOT

X -

B

D

— — ' _.„,-, u

0 ^S » O-- —

_

N»

-

POTASSIUM

1 1 1

iii i

7 -

6-

5-

.

1

1

1 1 1 1 1

-

0

•.

A \

\ *

\ ^

s^ b..

\\

LEGEND

lOAR FT FID PLOT

1936 FIELD PLOT

1937 FIELD PLOT

... B CD.

~" '^>^"^

-

SILICA- FREE ASH J

-

1

1

i i i i i

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 12. — March of silica-free ash and potassium in Avena harbata
for 1935, 1936, and 1937.

30

University of California — Experiment Station

w

DC

-

1 1

1

III 1

-

LEGEND

-

\A

1QA* FTFTD PLOT

_

1936 FIELD PLOT

o--

— -^ \

1937 FIELD PLOT

v \ \

-

■'••• \ \
•A \

-

-

V° \

\\ \

-

-

V- \
*\ \

POTASSIUM

-

\

•. \

-

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y

VAB

-

-

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.

-

©■-TT^SV^

^^;:;:^::::::^^s^

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_ I

-

i i

1

•©— —

III 1

EARLY OUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 13. — March of silica-free ash and potassium in Poa scabrella
for 1935, 1936, and 1937.

Bul. 627] Composition of Foothill Plants 31

nutritional value of the plant. Moreover, the mere analysis of a species
at a single growth stage affords little information as to the value of a
species as a range plant, nor does it contribute to the problem of mainte-
nance of the forage cover.

The potassium content in the seeds is low, being less than 1 per cent,
and is even lower than in the mature, weathered plant.

In calcium and phosphorus the rate of the seasonal decline for Avena
barbata and Poa scabrella is shown in figures 14 and 15. In calcium, the
rapid decline is from the early leaf stage to late bloom, designated as A B
in the figures, whereas in phosphorus there is a somewhat more gradual
and continuous decline from the time of early growth to maturity, desig-
nated as A C. In each of these constituents there is no appreciable change
in the levels from the time that the herbage dries to the end of the dry,
hot summer, designated as C D.

Shortening of the growing season of the grasses, since it affords only
a short period for absorption, evidently results in a lower level of those
constituents which occur limitedly in the soil solution, or those which are
absorbed slowly. The samples collected in the nursery represent plants
of a growing season shorter by more than 8 weeks than that of those col-
lected in the field plots, the seeding in the nursery having been done long
after growth had started on the range generally. At all points throughout
the growth cycle the percentages of calcium and phosphorus are much
lower than in the field samples, as typified in Avena barbata in figure 14.
In contrast to these lower levels, the length of the growing season has no
effect on the percentage of silica-free ash, potassium, protein, and the
other constituents studied, as seen in figures 8 and 12. The difference in
concentration and in the rate of supply of these three elements in the
soil solution may account for the difference in accumulation of these con-
stituents in the grasses studied. In a nearly neutral soil, such as that here
concerned, the phosphorus content is less than three parts per million.
In a short growing season the potassium content, being more abundant
in the soil solution than is phosphorus, for example, appears to be ab-
sorbed in much larger relative amounts. This finding may have an im-
portant bearing on the phosphorus content of grasses grown at high
elevations, or when limited precipitation curtails the length of the period
of growth. In January, 1937, on the range studied, when the tempera-
tures dropped so low as to freeze the tip of the young leafage, there was
a lowering of the phosphorus content in the early growth stage. This
low level was followed by rapid increase in phosphorus, as shown in seg-
ment A B of figure 14, after which the values of this constituent assumed
the usual trend.

32

University of California — Experiment Station

0.5

H

55

w

Uo.4

w

0.3

0.2

0.1

LEGEND

1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

CALCIUM

+■ C D

o

0.0
0.6

W

0,

0.2

LEGEND

1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

0.1 -

PHOSPHORUS

0.0

EARLY OUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig 14. — March of calcium and phosphorus in Avena barbata
for 1935, 1936, and 1937.

Bul. 627]

Composition of Foothill Plants

33

1.0

0.9

0.7

0.5

0.4

0.2

0.1

1

i i

111 l

-

LEGEND

\ A

1QAR FT ELD PLOT

\

1936 FIELD PLOT

\

1937 FIELD PLOT

\\

-

\\

\ v

\ \

\

-

\\

\\
. \\

CALCIUM

~

_

"•• A

\\

'•• \

_

•A

_

4b

_

-

%"■""•——-

^__^^cr^Tl c D "

~"*^— "Q-— «. ^° °" ° —

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0.5

0.3

0.2

1 1

1 1 1 1 1
LEGEND

1Q3R FTFTD PLOT

\A

1936 FIELD PLOT

V

1937 FIELD PLOT

X)

-

\

^"""l$v

..-■■•""t\ \

Or" \ \

\ \

PHOSPHORUS

\ \

\\

B

Y1--

V9"-^*"

"""^"-"S.:*-*

^^^^^

V"\ C D

N "••... •

—

X-r:r"

1 1

.©

i i i i .... . i

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 15. — March of calcium and phosphorus in Poa scabrella

34 University of California — Experiment Station

The tendency for the mineral constituents to change little in their
values after the blossoming period, as shown in segments C D of figure 14,
may be accounted for by decreased absorption due to curtailment of
physiological activities, associated, as this function is, with decreased
production of root hairs as the season advances.

The calcium content in Avena barbata, collected on the regular field
plot (table 1) shows a decline of 60.0 per cent in 1935 and 54.4 per cent
in 1937, from the first to the last collection. Such seasonal decline in cal-
cium holds proportionally for all the other species investigated. In the
same growth stage the various grasses studied show a variation in cal-
cium of approximately the same values as that for the entire growth
cycle of a single species. The following data illustrate this point for the
grasses in the early leaf stage, in 1937 :

Calcium,
Species per cent

Avena barbata 0.533

Bromus mollis 672

Bromus rigidus 922

Festuca megalura 402

Melica imperfecta 947

Boa scabrella 0.865

It is seen that a difference between the highest and lowest calcium content
of the six species is 51.2 per cent, compared with a difference of 54.4 per
cent in A. lariat a for the entire growth period.

In phosphorus content the same general tendency holds as for calcium,
with the exception that the decline after the blooming period is more
pronounced. From table 1 it may be calculated that the variation between
the highest and the lowest phosphorus content in Avena barbata for the
three years of study is 73.0 per cent, and in Bromus mollis it is 87.9 per
cent.10 This declining trend with advancement towards maturity holds
for all the other grass species. When, however, the same growth stage of
the various grass species is compared, the difference between them in this
constituent is relatively slight.

It is well known that the nutrients in the soil affect the composition
of vegetation. Table 1 contains data on forage produced on a heavily
manured plot located near the Station headquarters where the "A" hori-
zon is especially well developed. In Bromus rigidus, collected on this plot
in 1937, the phosphorus and protein contents are higher, whereas the

10 The figure for Avena barbata for 1937 is rather low for the growth stage con-
cerned, and is accounted for by the fact that the sample was collected from regenera-
tion growth following a heavy freeze of the earlier leafage. It will be noted from
table 1 that in later collections in 1937 the phosphorus content increases to average
amounts, and declines subsequently.

Bul. 627]

Composition of Foothill Plants

35

calcium and fiber contents are lower than those in the samples taken on
the regular field plot. The potassium content, on the other hand, is little
affected.

From these data, and unreported work of the authors, it would appear
that the soil solution and length of growing season modify the character-
istic levels of the constituents studied, except in the case of potassium
which is but slightly influenced.

The calcium content in the seeds is generally lower than that of the
mature herbage. The phosphorus content in the seeds is relatively high,

TABLE 2

Calcium-Phosphorus Ratio of the Grasses for All Growth Stages

for Each of Three Successive Years

1935
Field plot

1936

1937

Species

Field plot

Nursery
plot

Field plot

Avena barbata

1.0
0.7
1.2
1.2
1.0

1.2

1.0
1.1
1.2
1.2
1.0
1.3
1.5

0.9
1.2

1.6
1.6

1.0

Bromus mollis

1.0

Bromus riqidus

1.6

1.0

1.1

Melica imperfecta

1.8

1.5

and approximates the level found in the aerial growth up to the time that
the seeds begin to form.

The Calcium-Phosphorus Ratio. — The calcium-phosphorus ratio in
herbage is an important factor for foraging animals. To what extent this
ratio varies in the different groups of related plants is of scientific inter-
est and of economic importance. The ratios, for the grass species studied,
are presented in table 2.

Table 2 brings out the fact that in the grass species the averaged cal-
cium-phosphorus ratio for all growth stages is not far from 1. Although
the environmental factors may influence the percentages of calcium and
phosphorus, the data show that the effect is not such as to change the
ratio perceptibly.

Table 3 reports the calculated calcium-phosphorus ratio of Avena
barbata in seven growth stages. The data show that the calcium-phos-
phorus ratio in A. barbata holds near to 1 for all the growth stages. This
ratio is characteristic of the other grass species studied.

Evaluation of the Species as Forage. — From the preceding discussion
it is evident that all the grass species here investigated are high in caloric
values, and in minerals, from the early leaf stage until they have come

36

University of California — Experiment Station

into full bloom. Accordingly, the forage value of an annual range grass
is greatest during the interval of time between the earliest appearance
of the leaf blades and the date when the seeds begin to form, a period
when all essential constituents are maintained at a relatively high level.
Festuca megalura has a short growth period ; its value as range forage
is virtually over when the plant is in full bloom. From table 1 it is seen
that the late flowering period is reached in approximately 2 months after

TABLE 3

Calcium-Phosphorus Ratio of Avena Barbata at Various Growth Stages
for Three Successive Years

Growth .stage

1935
Field plot

1936

Field plot

Nursery
plot

1937
Field plot

Early leaf stage

Just before flowering

Plants in full bloom

Seeds in dough stage

Leaves mostly dry, some seeds cast. .

Aerial growth dry, seeds cast

Plants dry and weathered, seeds cast.

Average.

1.2*

0.8

0.9
0.8
0.9

0.9

1.0
0.9
0.9
1.0
1.4
1.3

1.1

1.0
0.8

0.9
0.8

1.5
0.9*
0.7

1.4
1.3
1.0

1.1

* Average of two samples at the same stage.

the earliest leafage appears. In Poa scabrella, although a perennial, the
interval between the early leaf stage and flowering is also short, and
the decline in protein and mineral constituents is rapid. Avena barbata
only slightly outranks F. megalura and P. scabrella as a range plant,
chemical characteristics alone considered. These three species are inferior
as forage because of the unfavorable nutritional balance at the onset of
blossoming. On the other hand, Bromus mollis, a species widely distrib-
uted and often abundant in the foothills of the state, contains a relatively
high percentage of protein at the time of blossoming ; and even at ma-
turity this constituent is relatively high because of the retention of the
seeds. The mature panicles, with the seeds intact, contain 12.53 per cent
crude protein and only 17.23 per cent crude fiber. B. rubens, although
mechanically objectionable as forage when the awns are developed, is
nevertheless of about the same nutritional value as B. mollis. Melica
imperfecta, a perennial which blossoms and matures late in the season,
maintains a high nutritional level even after the seeds have reached the
dough stage.

From the preceding discussion it becomes evident that the following
three characteristics are among the important factors in the evaluation

Bul. 627] Composition of Foothill Plants 37

of range grasses from the standpoint of their composition : (1) the in-
terval between the earliest appearance of leafage to full bloom must be
long; (2) the crude protein content must remain at a relatively high
level after the blossoming period, and well into the time that the seeds
attain the dough stage ; even after seed dissemination it is desirable that
a portion of the basal leaves and stems remain succulent until late in the
summer — a characteristic of many perennial grasses ; (3) in the species
whose herbage matures early, a large proportion of the seeds must be
retained after the leaf blades have become dry.

It is the opinion of the writers that the mineral constituents should not
in general be regarded as the deciding factor in the evaluation of the
range grasses studied. Indeed, the relatively few forage species which
contain unusually large quantities of sodium chloride, silica, and sele-
nium, are undesirable primarily because of the toxicity of the mineral
content. Ingle (26) and others have reported that forage plants contain-
ing as little as 0.05 per cent phosphorus are deficient for domestic live-
stock ; and Hall and Russell (20) concluded that 0.25 per cent phosphorus
is characteristic of superior forage.

All grass species here reported, and particularly those which retain
their seeds long after maturity of the herbage, are generally adequate in
calcium and phosphorus, provided these elements are fully utilized by
foraging animals. That components other than the percentage of the
mineral matter may determine the range value of a species (provided it
is palatable) is well illustrated in Avena harhata, which, although of
very low protein content when the seeds are cast, contains more silica-
free ash than any other grass species here investigated. Table 1 shows
that only in three instances, all of which occurred at full maturity in the
grass species studied, does the phosphorus decline to, or approach, the
deficiency level of 0.05 per cent. It is significant that these deficiencies
occurred in the grass samples collected in the Station nursery, where the
growing season was much shorter than on the regular field plot.

COMPOSITION OF GRASSLIKE SPECIES

Three grasslike species were included in the study, namely, Juncus bu-
fonius, J. oxymeris, and Eleocharis palustris. These species compose
much of the cover in the meadowlike swales of the San Joaquin Experi-
mental Range, and the latter two species furnish palatable forage in the
summer and early autumn after most of the true grass cover has become
dry (fig. 1) . This meadow type is generally under water for several weeks
during the period of heavy precipitation. Soon after the water recedes,
the stand forms nearly a complete ground cover, the herbage of which

38

University of California — Experiment Station

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Bul. 02 7] Composition of Foothill Plants 39

remains more or less succulent until late in August, and often into Sep-
tember.

The constituents of these grasslike species exhibit trends similar to
those of the corresponding components in the grasses. There is an orderly
trend in the same direction in the organic and inorganic constituents
from the early leaf stage to maturity. The change in composition is rapid
from the time of the earliest appearance of leafage to the flowering stage.
From then on the changes are slight, if they occur at all.

Organic Constituents. — Table 4 shows that there is little difference in
the crude protein content in the three grasslike species here reported,
when the same growth stages are compared. When the entire growth
cycle of the species is considered, however, there is a decline in the pro-
tein content from the early leaf stage until the herbage is dry.

The crude fiber content in the three species studied shows the same
general behavior, being lowest in the early growth stage and highest at
maturity. After the flowering period is reached there is little increase in
fiber.11 In J uncus oxymeris, for example, the fiber content is 50.1 per cent
higher in the flowering stage than in the early leaf stage and changes
but little to full maturity — leaf stage — these comparisons being made
from 1936 and 1937 data. The data show that the difference in the per-
centage of protein and of fiber for a single growth stage of this plant
group is less than that for any one of these species for the entire period
of growth.

Inorganic Constituents. — Table 4 shows that Eleocharis palustris is
conspicuously high in silica, containing, in fact, the highest percentage
of this constituent of any species investigated, reaching 12.33 per cent
at maturity.12

The silica-free ash in the grasslike species is highest in the early leaf
stage. After reaching the bloom stage there is little change in this con-
stituent. The levels of ash of the species of this group are similar at
specific growth stages.

In calcium content these species lack consistency in trend from the
early leaf stage to the flowering period. From the latter period on to
maturity, however, there is little fluctuation. In phosphorus content, on
the other hand, there is a gradual decrease from the early leaf develop-

11 The slight irregularities in the trend of the fiber representing the various growth
stages is presumably accounted for by the fact that the samples, though pure as to
species, were necessarily obtained from the average growth of a dense stand, rather
than from selected individual specimens.

12 The high silica content at all growth stages of this species is significant in view
of the fact that the samples immediately after collection were washed in several
changes of water ; but it is possible that the surficial soil silica was not entirely re-
moved from the floral parts.

40 University of California — Experiment Station

ment to maturity, reaching the lowest point when the leafage has dried.
In potassium content little change takes place through the life cycle, be-
ing only slightly higher in the early leaf period. J uncus bufonius is the
lowest in potassium and silica-free ash of the three species.

Evaluation of the Species as Forage. — In these species the calcium-
phosphorus ratio is approximately 1 in early stages of development,
whereas at maturity it approaches 4. As stated, the primary value of
most of these species as range forage is that their succulence remains
high when the herbage of most of the grasses and the broad-leaved species
has dried. Although the protein values are relatively low, and the fiber
content fairly high in midsummer, the levels of all the nutrients at that
season are much higher than in the other herbaceous species. Eleocharis
palustris, being very high in silica, appears to be of lesser value than the
other two species. In silica-free ash and its constituents the grasslike spe-
cies rank favorably with those of the grasses during the entire period of
growth.

COMPOSITION OF THE BROAD-LEAVED HERBS

Twenty-five broad-leaved herbs were included in the study, these being
representative of seven plant families. The list includes the most impor-
tant forage species, and also some "range weeds," or plants of very low
palatability for domestic foraging animals. The weed species studied are
of interest because of their occasional conspicuous abundance in the
plant community, and their presumed effect upon the soil and on plant
succession.

Organic Constituents. — Figure 16 shows the trends of crude protein
and crude fiber in Amsinckia Douglasiana — this species typifying fairly
well the behavior of these constituents for the broad-leaved herbs. An
impressive consistency exists in the levels of the protein and fiber con-
tents in the same growth stage for the three successive years of study.
The rate of change in protein and fiber is most rapid from the early leaf
stage to the late blossom period, designated as A B in figure 16. From
the latter period to maturity, indicated as B C, the change is less pre-
cipitous. The relative rate of change of these two growth stages holds for
all but one group of species of the broad-leaved herbs studied. In the
species of the legume family, as represented by Lupinus Benthami, the
protein content invariably declines slowly and at nearly the same rate
throughout the life cycle. It should be noted that the maximum and mini-
mum protein and fiber levels, as recorded early in the season and at
maturity, respectively, are of slightly higher values in the legumes than
in the nonleguminous species. In the former species, the protein level

Bul. 627] Composition of Foothill Plants 41

remains relatively high throughout most of the growth period. It will
be noted (table 5) that the characteristic protein values of the broad-
leaved species are little or not at all affected by slight variations in the
length of the normal growth cycle. In the nursery, where the growing
season was short, the percentage of protein for a given growth stage is
of the same absolute value, and the rate of the seasonal decline, as shown
by the shape of the curve (fig. 16), is identical with that of samples col-
lected in the regular field plot.

Table 5, which gives the analytical data of this group, has similar ar-
rangement to that of tables 1 and 4 ; but in this table the species are
grouped consecutively according to family relationships. In stating the
particular growth stage which the various samples represent, it should
be recognized that not only is there variation in the growth stage of the
individual plants which composed the sample, but also that parts of in-
dividual plants differed somewhat in the stage of development. Species
of Erodium and Amsinckia are particularly difficult to sample for the
reason that a single plant may simultaneously display a combination of
buds, newly expanded flowers, and well-formed seeds. Although the vari-
ous samples are composed predominantly of the growth stage designated,
some material of a different stage of development may inadvertently
have been included.

Table 5 shows that the protein content, although differing widely in
amount in the various species, is highest in the early leaf stage and lowest
at plant maturity. During the three years of investigation the range in
crude protein for all species worked with, when in the early leaf stage,
is from 16.07 per cent to 35.05 per cent, and at maturity, when the seeds
have been cast, the range is from 12.36 per cent to 1.38 per cent. The
variation within a species is fairly well typified in Silene gallica, in which
there is a decline of 76.6 per cent during the growth cycle of 1936. In con-
trast with this wide descending seasonal trend within a species the range
in protein content in the early growth stage and at maturity is relatively
slight. This point is shown in table 6, compiled from table 5, which in-
cludes species belonging to different families.

The levels in protein content among the four species, in the early
growth stage and at maturity,13 respectively, are seen to vary but little.
Moreover, the percentages of protein differ little from year to year in a
given species. These species, however, show a characteristically wide vari-
ation in the protein levels at the flowering period, as seen when Lupinus
Benthami and Hemizonia virgata are compared.

13 The relatively high protein content reported for Lupinus Benthaim is accounted
for by the fact that portions of the pods retained their seeds at this sampling period.

42

University of California — Experiment Station

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52

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Bul. 627]

Composition of Foothill Plants

53

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54

University of California — Experiment Station

The seeds of this plant group are much richer in crude protein than
are those of the grasses. This is especially true of Lupinus Benthami and
Lotus americanus, whose seeds have a protein content in excess of 46
per cent.

In crude fiber the broad-leaved species show the expected behavior of
relatively low content in the early growth period, and of maximum
amounts at plant maturity. "When the plant is young the percentage of
growing tissue is relatively high and the cell walls are in a formative

TABLE 6

Variation in Percentage of Crude Protein in Four Species

of Broad-leaved Herbs for Three Successive Years*

Early leaf stage

Flowering stage

Plant maturity

Species

1935

1936

1937

1935

1936

1937

1935

1936

1937

Amsinckia

Douglasiana

23.95

22.71

23.43

10.63

13.85

10.61

4.20

4.29

4.95

Erodium botrys

30.35

24.95

19.94f

18.66

15.11

4.61

4.10

4.17

Hemizonia virgata .

26.20

22.94

21.61

6.59

6 49

6.92

5.30

5.32

Lupinus Bentha m i

27.61

27.55

22.06

21.15

17.26

6.75

4.63

8.71

* In cases where more than one anah Ufl was made of a given growth stage, the results were averaged.
t This low value may he accounted for by the fact that the first aerial growth of the season was greatly
slowed down for ahout a month by the abnormally low temperatures.

stage. At maturity the cell walls are Lignified and the proportion of cel-
lulose is high ; this accounts for the high fiber content. In some species
like Filago gallica, however, which build up high fiber content early in
the growth cycle I he stems consl h ate a Large pari of the total weight of
the plant. In the same stage of growth for the three successive years of
study the fiber content of a given species is much the same, indicating
slight response of this consl ii aenl to environmental variations (table 5).
The range in crude liber through the entire growth period is wide for
any one species of this group, but it is relal ively narrow in a given growth
stage when the various species are compared.

In the seeds, the crude fiber content of this plant group ranges from
approximately 7 to 19 per cent. In the seeds of the leguminous species,
however, the fiber content is very Low.

Inorganic Constituents. — The rate of decline in the percentage of
silica-free ash, and in potassium content, with respect to seasonal devel-
ment is typified in Amsinckia Douglasiana. The decline in these constitu-
ents from the early Leaf stage to the late flowering period is rapid, as
shown in segmenl A B of figure 17. From the late flowering period to
maturity the level of the ash changes little, as seen in B C of the figure.
A single analysis made at any point along the segment A B is seen to be

Bul, 627]

Composition of Foothill Plants

55

35

25

U5Q_

15

25

1 1

1

III 1 _

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FIBER

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/ /

LEGEND

~

/ /

/ /

IC^ft ^IID^FPV

/ /

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1Q.^ FTF.I.n PLOT

1936 FIELD PLOT

- ' </

1937 FIELD PLOT

6

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1 1

1

III 1

20 -'

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1

1 1

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LEGEND

: a

- \ v°

IQ^A WTTDSTTOV

1935 FIELD PLOT
1936 FIELD PLOT

- x^\

1937 FIELD PLOT

-© \x\

-

-

PROTEIN

-

1\ B

-

.

>•.

-

<*■ — -^^^^^ — ~--o

1

1 1

iii i

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 16. — March of crude protein and crude fiber in Amsinckia Douglasiana
for 1935, 1936, and 1937.

56

University of California — Experiment Station

1 1 1

w
u

a

Q* 3

LEGEND

__ 1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

POTASSIUM

i r

20

\8

U

a

CL12

10 -

LEGEND

1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

1 -

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 17. — March of silica-frco ash and potassium in AmsincMa Douglasiana
for 1935, 1936, and 1937.

Buk 627] Composition of Foothill Plants 57

of little value in signifying the levels of these constituents for the early
growth stages. On the other hand, a single analysis representing a sample
characteristic of the segment represented by B C, gives a fairly reliable
criterion of the ash and the potassium content of the plant from the
flowering period to maturity.

There is wide variation among the percentages of silica-free ash in the
various broad-leaved species at a given growth stage. Some species are
consistently high in ash, whereas others are low in this constituent in
the same developmental stage. Consistently high percentages of silica-
free ash are found in the species of the borage family, among which
Amsinckia Douglasiana and Plagiobothrys nothofulvus contain the high-
est amounts, the ash content accounting for nearly one-fifth of the mois-
ture-free weight of these species when in the early leaf stage.

Although the levels of the silica-free ash vary among the different
species, the percentages within a given species show still greater varia-
tion during the growth cycle. Thus in Lupinus Benthami and Hemi-
zonia virgata, in 1936, the ash content declines 41.4 per cent and 69.0 per
cent, respectively, from the earliest leaf stage to maturity. In Er odium
cicutarium, on the other hand, the variation is relatively slight, being
14.49 per cent in the early leaf stage in 1935, and 11.00 per cent at ma-
turity in 1936. Of all the broad-leaved herbs investigated, the three spe-
cies of Er odium show the least variation in ash content during the entire
growing period.

Table 5 shows much consistency in the silica-free ash content within a
species for any single growth stage, during the three years of study. Wide
variation in the silica-free ash is recorded for the various species at the
time of blossoming. The species which blossom early in the season, such
as Er odium botrys, E. cicutarium, Lupinus bicolor, and Plagiobothrys
nothofulvus, are high in silica-free ash at that stage ; those which blossom
late, such as Hemizonia virgata, Lessingia germanorum, and Lotus amer-
icanus, are low in ash content when in the flowering period. Accordingly,
late-blossoming species appear to be characterized by lower ash content
in the later stages of vegetative growth.

The ash content in the seeds of the broad-leaved herbs is of a slightly
lower level than that of the dry vegetative portion of the plant collected
on the same date. The seeds of the leguminous species are the lowest in
ash of this entire plant group, whereas those of Amsinckia Douglasiana,
the three species of Erodium, and Plagiobothrys nothofulvus are the
highest in this component.

The potassium content accounts for more than one-third of the silica-
free ash in all the broad-leaved herbs. The maximum percentages are

58 University of California — Experiment Station

invariably found in the early leaf stage. In all species the levels of this
element decrease from the early growth period to plant maturity. If the
young seedling is regarded as the beginning of plant development, and
maturity of the seed as the culmination of the life cycle, then the level
of potassium decreases from initial growth and reaches the minimum in
the fully developed seed.

In the early leaf stage Amsinckiana Douglasiana and Plagiobothrys
nothofulvus contain the highest potassium content recorded in these
studies. These are followed by Hemizonia virgata and Gilia tricolor, re-
spectively, in the list of potassium-rich plants. The lowest potassium
content recorded in the early leaf stage is in Lotus americanus, with 2.50
per cent, and in Lupinus formosus, with 2.51 per cent. From the early
growth stage to full plant maturity the decline in potassium is somewhat
more precipitous in species especially rich in this element than in those
characterized by low potassium levels in the early leaf stage. In Plagio-
bothrys nothofulvus in 1936 there is a seasonal decline of 81.9 per cent. In
contrast, in species of lower potassium content, such as Erodium botrys,
there is a decline of 60 A per cent through the season.

In the seeds the potassium content is low ; in fact it is lower than in
that of the mature, dry herbage, ;i reversal as compared with the phos-
phorus content. Thus in the seeds of Lupinus Benthami there is 1.94 per
cent potassium, whereas the Lowest content in the vegetative parts for the
three years of investigation is 2.10 per cent. In the seeds of Amsinckia
Douglasiana there is 1.35 per cent potassium, whereas in the vegetative
portion there is 2.4.") per cent, this figure being the lowest recorded for
this species.

The data show thai there are conspicuously rich, as well as compara-
tively poor potassium plants among the broad-leaved species investi-
gated. Since all of these species vary widely in potassium content through
the growth cycle, recognition of their potassium level can only be known
when analyses arc made ;it several growth stages.

In calcium content tin' composition remains at nearly the same level
throughout the life cycle in mosl of I he broad-leaved species sampled in
the regular field plots. Figure 18, which gives the seasonal trend of the
calcium content in Amsinckia Douglasiana, shows that virtually the same
level of this constituent is maintained from early leaf stage to maturity.

Shortening of the growing season appears to affect little the calcium
content of the broad-leaved group of plants, as seen from samples pro-
cured in the nursery-phmtcd species, which were seeded not less than 2
months after growth had started on the regular field plots. This point
is typified in figure 18, where the percentages in calcium content for

Bul. 627]

Composition of Foothill Plants

59

2.0

1 1 1

A R

- A / \

«& \ • S.

1 1 1

i
C D.

o

-

CALCIUM

-

-

LEGEND

,_ .. IQ^fi >ITTRQF.DV

"

1935 FIELD PLOT
1936 FIELD PLOT

1937 FIELD PLOT

1 1 1

i i i

1

1.2

0.4

0.2

0.0

I I

LEGEND

1936 NURSERY

1935 FIELD PLOT

1936 FIELD PLOT

1937 FIELD PLOT

PHOSPHORUS

EARLY JUST FULL SEEDS IN SEEDS PLANTS PLANTS DRY PLANTS DRY

LEAF BEFORE BLOOM. DOUGH MATURE MOSTLY AND

FLOWERING STAGE DRY WEATHERED

Fig. 18. — March of calcium and phosphorus in AmsincMa Douglasiana
for 1935, 1936, and 1937.

60 University of California — Experiment Station

Amsinckia Douglasiana, of the nursery- grown samples, are seen to differ
but slightly in their values as compared with the field samples. Table 5
shows this tendency also to hold for the other nursery-grown broad-
leaved herbs. Samples representative of species grown near the Range
Station headquarters on a deeper soil are of about the same calcium con-
tent as those collected on the regular field plot.

In phosphorus content the trend of the various species studied is repre-
sented by Amsinckia Douglasiana (fig. 18). There is a somewhat more
rapid decrease in phosphorus from the earl}7 leaf stage until the late
flowering period, designated as A B on this graph. From the flowering-
stage to maturity the decline continues, but at a less rapid rate. The phos-
phorus content in the nursery samples, as illustrated in figure 18, main-
tains the same general trend as in the regular field samples, but the
absolute values are much lower. The apparent adverse effect of the short-
ened growing season on the levels of phosphorus holds for all nursery-
grown species.

The phosphorus content in the broad-leaved species maintains the most
orderly behavior of the minerals investigated. The minimum percentage
is reached when the plants are dry and the seeds cast. The highest phos-
phorus content recorded for any growth stage of the broad-leaved species
studied is 1.130 per cent, found in OUia tricolor; whereas the lowest
amount obtained is 0.040 per cent, in Trifolium tridentatum. When the
extreme range in phosphorus content for a given growth stage is consid-
ered, for example the flowering period, ihe Largesl amount is 0.724 per
cent, recorded for Erodium cicutarium, and the smallest amount is 0.128
per cent, recorded for //< mizonia virgata.

The members of the Legume family, as seen in table 5, are the lowest in
phosphorus eontenl of the broad-leaved herbs. This constituent is espe-
cially low at plant maturity for cadi of i lie six leguminous species. These
figures bring out the fact that the same general behavior in the phos-
phorus content holds for the broad-leaved plant "/roup, as in the grasses
and grasslike species; namely, that the variation in phosphorus in a given
species through the entire growing season is much greater than that in
the various species for a specified growth stage. Obviously, therefore,
determination of the phosphorus content only once or twice during the
growing season may be of little value in evaluating the levels of this con-
stituent in a range plant.

It is noteworthy that in nearly all the species the seeds contain a high
percentage of phosphorus, and that the levels compare favorably with
those of the very young leafage. In the seed of Plagiobothrys nothofulvus,
for example, the phosphorus content is 0.842 per cent, and in the very

Bul. 627]

Composition of Foothill Plants

61

young leafage it is 0.759 and 0.843 per cent. In Amsinckia Douglasiana
there is a range between 0.766 and 0.878 per cent in the seed and 1.080 in
the early leafage in 1935. If the seed is regarded as the initial stage of
plant development, the percentage of phosphorus decreases from the very
initiation of life-cycle activities, beginning with seed germination, and
continues downward until full maturity is reached.

The Calcium-Phosphorus Ratio. — In the broad-leaved herbs it has been
shown that the calcium content remains at almost the same level through

TABLE 7

The Calcium-Phosphorus Ratio of Broad-leaved Herbs in
the Early Growth Stage and at Maturity*

1935

1936

1937

Species

Early
leaf
stage

Maturity

Early
leaf
stage

Maturity

Early
leaf
stage

Maturity

1.6
2.0

2.9

1.0
2.6

2.9

2.1

2.2

6.8

8.1
6.1
3.5
15.3

6.2

7.7

3.5

2.8
7.6

3.7
2.3

4.6

2.5
2.6

3.7

8.1

4.9

15.0

27.9

24.7

12.7

8.5

6.8

13.2

12.5

13.0

7.6

8.1

12.5

3.4

2.7
5.6
8.8
6.5

4.1

4.7
3.7
3.9
2.5

4.6

6.2

8.5

30.4

55.6

15.8

3.8

4.3

6.8

5.7

15.2

* In cases where more than one analysis was made of a given growth stage, the results were averaged.

the entire growth cycle, whereas the phosphorus content declines stead-
ily, to reach the lowest level at plant maturity. This behavior results in a
continuous widening in the calcium-phosphorus ratio with advancement
of the season.

Table 7, compiled from table 5, shows that the calcium-phosphorus
ratio is lowest in the early leaf stage, with an average range for the sev-
eral species of 2.2 to 4.6 for samples taken on the regular field plots dur-
ing the three years of study. At plant maturity the corresponding ratios
are 7.7 and 15.2. This wide ratio at maturity is accounted for chiefly by
the low phosphorus content. The calcium-phosphorus ratio is seen to vary
widely in the species studied regardless of the stage of development.

Evaluation of the Species. — The broad-leaved species as a group, chem-
ical composition alone considered, fall into two categories as applied to

62 University of California — Experiment Station

the economy of the range : (1) species having a short vegetative period,
but which maintain a high level of protein and ash contents well into the
late flowering stage ; (2) those having a long vegetative period and char-
acterized by a low level of ash and protein when the flowering period is
reached. Erodium botrys affords an example of the first of these cate-
gories, and Hemizonia virgata, of the second group.

In Erodium botrys, from the early leaf stage to flowering, an interval
of 4 weeks (from February 8 to March 6, 1936) , the decline in crude pro-
tein is only 20 per cent. But from the flowering stage to maturity of the
herbage, a period of 9 weeks, this constituent decreases nearly 75 per
cent. In contrast, there is a decline in the late-blossoming Hemizonia
virgata of 77 per cent from the early leaf stage to the time of flowering,
a period of 27 weeks (March 6 to September 10, 1936), and of only 18
per cent from the flowering period to maturity, an interval of 8 weeks.

Table 5 shows that the ash, and some of its constituents, maintain ap-
proximately the same trends as do the protein. The relatively high nutri-
ent content during a long vegetative period, however, enhances the value
of palatable plants having late-flowering characteristics, despite the low
values of the important constituents when the flowers appear.

The high protein content reported throughout the growth cycle for
the palatable leguminous species, despite the short period between early
growth and flowering, would appear to place the plants of this genus in
a very favorable economic position.

The phosphorus content of this group, at plant maturity, however, is
among the lowest of any of the herbs whose samples were collected in the
regular field plots, the percentages being in the second decimal. It is
noteworthy that of all the leguminous species studied, Lotus americanus
and Lotus strigosus — the former very common on the foothill ranges —
are the lowest in ash and its various constituents at all the growth stages.
Despite these apparent deficiencies in inorganic constituents, their rela-
tively high yield of protein throughout the season places them among the
most valuable of forage species as far as nutrients are concerned.

Of the species of the borage family, Amsinckia Douglasiana, Plagio-
bothrys nothofulvus, and Plagiobothrys tenellus, all of which are early
maturing, maintain a high level of protein well into the late flowering
stage, and of ash and its constituents until maturity. An exception is
seen, however, as pointed out, in the collections taken from the late-
planted nursery plots, where the phosphorus content of the samples is
low. Cryptantha flaccida, a species which has a long growing season, and
which occurs commonly on depleted, thin soils in the locality studied, has
the lowest percentage of protein and inorganic constituents at all growth

Bul. 627] Composition of Foothill Plants 63

stages of the four species of this family. Although most of these boragi-
naceous species here concerned are not of especially high palatability,
they are generally high in nutritive values.

Among representatives of the geranium family, all of which mature
early, Erodium cicut avium, as previously stated, has the highest content
of protein and silica-free ash of the three species studied. Moreover, the
high levels of all the constituents studied in this species are fairly well
maintained to full maturity. Comparison of the three species when fully
mature, as in 1936, shows a range in protein content of from 9.98 to 12.36
per cent for Erodium cicut avium, of from 3.36 to 4.84 per cent for E.
botvys, and of 4.36 per cent for E. moschatum. The phosphorus content
also is much the highest in Ev odium cicut avium.

Table 5 shows that Ev odium botvys and E. moschatum have nearly the
same nutritive values. Collections of a divergent form of Erodium botvys,
grown near the Station headquarters, in which a single plant composed
each sample, were analyzed for comparison with material procured on
the regular field plot. This divergent form is more robust, later maturing,
has larger flowers, longer styles, and the inner sepals have a conspicuous
red margin. Table 5 shows this form to be lower in all constituents, except
in fiber, than the more typical forms of Erodium botvys. Possibly the
form in question justifies its recognition as a variety.

Of the four species of the composite family, the late-maturing
Hemizonia vivgata, although of low palatability, is nutritionally superior
because of the high levels of protein and minerals throughout the long
vegetative period. Filago gallica and Lessingia gevmanovum, being low
in protein and ash constituents, must be regarded as distinctly inferior
species. Layia pentachaeta, although having a short vegetative span, at
no time maintains a high level of the essential constituents. Even when
very young, it approaches in composition the other inferior species of
this family. The three above-named species are low in protein and ash
at the growth periods when the other early-maturing species are still high
in these constituents.

Gilia tvicolov, another early-maturing plant, is characterized by high
composition of protein, ash, phosphorus, and potassium. At maturity all
constituents recorded, except fiber, reach relatively low levels. Navav-
vetia viscidula, also a member of the gilia family, is similar in composition
to that of Gilia tvicolov, except that the phosphorus content is lower.

Godetia viminea and Godetia amoena, of the primrose family, are con-
spicuously low in all important constituents investigated except calcium.
At maturity both species are very low in protein and ash.

Silene gallica, the only species studied of the pink family, is relatively

64 University of California — Experiment Station

low in protein in all growth stages, as compared with the other broad-
leaved species, but is fairly high in ash, with a satisfactory balance in
essential mineral constituents.

COMPOSITION OF BROWSE FROM SHRUBS AND TREES

Since the foothill area investigated is of the open savanna-woodland type,
browse vegetation contributes to the forage supply of the range at certain
seasons, and presents a problem somewhat distinct from that of the life
forms previously discussed. The nine species studied include deciduous
and nondeciduous (evergreen) shrubs and trees of wide distribution.
Because most of these species either mature late in the season or are
green throughout the year, they supply important nutrients when the
herbaceous vegetation is dry.

Table 8 gives the analytical data of the samples collected at various
growth stages. The data are discussed separately for plants of the "half-
shrub" type of the deciduous trees and shrubs, and of the nondeciduous
shrubs.

Half-shrubs. — The samples of the two species of half-shrubs, charac-
terized by semiherbaceous stems, namely, Lotus scoparius and Lupinus
albifrons, contain both leaves and si cms of the current growth. Since the
proportion of these parts unavoidably varied in the different samples
collected, i he cyclic I rends of I lie composil ton are somewhal irregular.

The crude protein content in the shoots of these two leguminous species
decreases, and the crude fiber increases, from the early leaf stage to ma-
turity. In Lupinus albifrons, however, the protein Levels are consistently
higher throughout the season, whereas the current growth of Lotus sco-
parius contains a lesser amount of crude fiber earl}r in the season. The
ash content, and the potassium and phosphorus of I aese species, decrease
from the early leaf stage to maturity, whereas the calcium content shows
no definite trend. Potassium comprises the highest percentage of the ash,
followed by calcium and phosphorus respectively.

The leafage of the true woody deciduous shrubs and the trees — species
wdiose stems are not green or herblike — differ from the species previously
discussed, as seen in table 8, in the march of composition of several of
the constituents. In this group the samples collected were of the same
age and were collected approximately the same distance from the ground.

The protein levels decrease with maturity, whereas the fiber content
follows no definite trend and is comparatively low. The ash and calcium
contents increase uniformly from the early leaf stage to maturity,
whereas the potassium and phosphorus contents decrease during this
period.

Bul. 627] Composition of Foothill Plants 65

Compared with the other plant groups discussed, the protein content
of the foliage in this group is very high in the early leaf stage, reaching
nearly 40 per cent in some species. Even when the foliage is mature, the
protein content is higher than in most of the species studied.

Table 8 shows that in the newly expanded flowers of Aesculus cali-
fornica and Sambucus glauca, including the pedicels, the protein content
is of the same level as that of the foliage for the corresponding period.
As the flowers fade and the petals are dropping, however, the protein
decreases much more rapidly than in the leaves.

The fruits of Aesculus calif ornica, when in the dough stage, are nearly
identical in protein content with the leaves collected on the same date,
whereas this constituent in the fruits of Sambucus glauca is of much
lower level than that in the leaves collected at the same time. The fruits
(acorns) of Quercus Douglasii are conspicuously low in protein, a point
of significance in view of their extensive utilization by foraging animals.
The mature, hulled fruits contain only about 5 per cent protein, whereas
the hulls, or shells, contain about 3 per cent — these figures being much
the lowest of any of the fruits studied.

The crude fiber content of the leaves of the deciduous shrubs and trees
is low, as stated ; indeed, it varies little during the entire growth cycle.
In Quercus Douglasii the maximum value in fiber for the three years of
study is 17.50 per cent and the minimum, 11.50 ; whereas in Rhus diver-
siloba the values are 14.55 and 12.89 per cent. The flowers in Sambucus
glauca are also low in fiber content, exhibiting much the same level as the
leaves, whereas in the fruits this constituent is generally slightly higher
than in the foliage. As would be expected, the hulls of the fruit of
Quercus Douglasii have a high fiber content, whereas the naked, or hulled
fruit, contains only from 4 to 6 per cent fiber. The young fruit of Aescu-
lus calif ornica, unhulled, contains 12 per cent.

The ash content in the foliage of the species of the true shrub-and-tree
group exhibits a reverse ratio to that of most species previously discussed,
being highest when the leaves are fully mature, and lowest in the early
leaf stage. For example, in 1936, the silica-free ash content in Aesculus
calif ornica increases 178 per cent with the progression of the season,
whereas in Sambucus glauca there is an increase of 65 per cent. The other
species of this group show much the same relation.

The calcium content forms the largest single fraction of the ash con-
stituents in the deciduous, woody plant group. There is an orderly in-
crease in calcium throughout the leaf developmental period. In some
species the percentage is exceedingly high at maturity of the foliage. In
Aesculus calif ornica, for example, the calcium increases more than 1,000

66

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72 University of California — Experiment Station

per cent from the time of unfolding of the leaves until they are mature
and dry. In Sambucus glauca and Quercus Douglasii in 1936 there is an
increase of 172 per cent and 151 per cent, respectively, for the corre-
sponding period. In the flowers and the fruits the calcium content corre-
sponds closely in values to that of the very young leafage.

The phosphorus content in the foliage is in reverse ratio to that of the
calcium, being highest in the young leaf stage and lowest when the leaves
are mature. Thus in Aesculus calif ornica, in 1936, the phosphorus de-
creases 67 per cent during the growing season ; in Quercus Douglasii and
Sambucus glauca it decreases about 64 per cent and 90 per cent, respec-
tively. In the young flowers the phosphorus content corresponds closely
to that of the very young leafage ; but as the flowers begin to fade this
element assumes approximately the same level as that in the foliage col-
lected on the same date.

The potassium content in the foliage of this plant group varies widely.
Quercus Douglasii is notably low in this element, whereas in Aesculus
calif ornica, Sambucus glauca, and Rhus diversiloba, the potassium con-
tent is high. In all four species this constituent is much the highest in the
early leaf stage, and declines in an orderly way to leaf maturity. The
decrease, in 1936, amounts to about 30 per cent in Aesculus calif ornica,
66 per cent in Rhus diversioloba, and 59 per cent in Quercus Douglasii.
The young stems of Sambucus glauca, as seen in the samples collected in
1936, are higher in potassium than the foliage for the corresponding col-
lection. The flowers and fruit of the species studied have approximately
the same level of potassium as the foliage for the corresponding growth
period.

N on deciduous Species. — In the nondeciduous species the new leaves
are interspersed with those produced the previous season. The composi-
tion of the samples here reported tends to vary but slightly throughout
the growth cycle, because of the admixture of new and old leaves.

Table 8 brings out the fact thai the crude protein content of the foliage
of Ceanothus cuneatus and of C. divaricatus is low, ranging from about
17 per cent to 8 per cent. In both species the highest protein content is
recorded when the proportion of young leafage is higher than the old
in the sample. It is significant that the protein content of the foliage of
the two species of Ceanothus is considerably higher from midsummer
until late in the autumn than in that of most of the mature herbaceous
species.

In the flowers of the two Ceanothus species protein is relatively high,
whereas in the fruits it is low, being of the same magnitude as that of the
leafage at the corresponding period.

Bul. 627] Composition of Foothill Plants 73

The crude fiber in the Ceanothus species is among the lowest of any
plants here reported, ranging from 5.44 per cent to 12.52 per cent. The
flowers contain nearly the same percentage of fiber as the leaves. In the
mature fruit, however, this constituent is relatively high.

The silica-free ash content in the three nondeciduous shrubs is rela-
tively low, Ceanothus divaricatus on the average being the lowest of this
group, and Bhamnus calif ornica the highest. No definite trend is shown
in the ash content in these species through the season, presumably be-
cause of the mixed character of the samples, as explained.

The foliage of these shrubs contains more calcium than any other single
mineral constituent. In the flowers and fruits of the Ceanothus species,
however, the percentage of calcium clearly ranks second to that of potas-
sium (table 8).

In potassium content Bhamnus calif ornica is the highest and Ceano-
thus cuneatus is the lowest of the three nondeciduous species. In most of
the samples of the Ceanothus species the percentage of potassium is in
the first decimal, whereas in no instance does the calcium content fall
below 1 per cent in the three species.

In phosphorus content the nondeciduous species, while not high at any
stage of growth, is maintained at a fairly constant level. For the two
species of Ceanothus the range for the three years of investigation is from
0.089 to 0.299 per cent. The flowers are richer in phosphorus than in any
other plant organ reported.

The Calcium-Phosphorus Batio. — In the deciduous shrubs and trees
the calcium-phosphorus ratio increases with advancement of the season,
being lowest in very young foliage and highest when the leaves are fully
mature. The ratios for 1936 are shown in table 9, as derived from table 8.

The data in table 9 are impressive of the fact that the calcium-phos-
phorus ratio is much the lowest in the early leaf stage and very high at
leaf maturity. In Aesculus calif ornica and Quercus Douglasii the rise in
this ratio takes place largely after the flowering period. In the young
leafage, when the meristematic tissue is proportionately large, and when
cell division is actively taking place, the percentage of phosphorus is
relatively high. But with advancement of age, and decline in the rate of
growth, the percentage of phosphorus decreases in the foliar organs.
Calcium, on the other hand, enters limitedly into meristematic activities,
but is utilized much more extensively for cementing the cell walls in the
older leaves with calcium pectate. Calcium also serves to neutralize the
by-products of metabolism, hence the percentage of its content reaches
the maximum when the foliage is mature.

In the nondeciduous group the calcium-phosphorus ratio varies greatly

74

University of California — Experiment Station

through the year. In Ceanothus cuneatus the ratio ranges between 6.1
in March, 1936, when the new leafage is partly formed, and reaches 19.7
in September, when the seeds are cast. In C. divaricatus the ratio ranges
from 4.2 in April, when in the flowering stage, to 15.1 in January, 1937,
before the new leaves are formed. In Bhamnus calif ornica the ratio
ranges from 5.6 in May, 1937, in the early fruiting stage, to 17.1 in Sep-
tember, 1936, when growth is at a minimum.

In the deciduous half -shrubs the calcium-phosphorus ratio likewise is
highest at the end of the growing season. In Lupinus albifrons, in 1936

TABLE 9

The Calcium-Phosphorus Ratio of Foliage of the Deciduous Shrubs

and Trees at Three Growth Stages ; 1936

Species

Early leaf
stage

Full bloom

Maturity

0.7
2.0
0.4
2.2

4.2
2.5
3.4

23.9

50.7

19.5

15.4

in the field plot, this ratio ranges from 3.4 to 8.7. In Lotus scoparius the
highest ratio is 7.6, recorded when the leaves are cast in September, 1936 ;
and it is the lowest, namely 2.7, before early bloom, in March of the same
year.

Evaluation of the Species. — The woody species discussed fall into two
categories in the matter of nutritional characteristics of the foliage,
namely, those whose chemical values change but relatively little during
the growing season, and those whose levels of the more important con-
stituents change markedly during the growth cycle.

The nondeciduous shrubs, namely, Ceanothus cuneatus, C. divaricatus,
and Bhamnus calif ornica, typify the species of the first category. Al-
though the levels of the organic and inorganic constituents of the foliage
in these species are relatively low at all times, they are maintained
throughout the year at more or less the same values. Because of the lower
crude-fiber content, and the favorable balance of the more important
nutrients, C. divaricatus outranks nutritionally C. cuneatus. B. calif or-
nica, on the other hand, being somewhat higher in silica-free ash and in
phosphorus than the two species of Ceanothus, appears to be the most
desirable. Despite the fact that these three species are at no time espe-
cially high in protein, and in total ash and its constituents, they are
nevertheless important on the range because of their succulence through-
out the year.

Bul. 627] Composition of Foothill Plants 75

The species of the second group — those whose chemical values change
markedly through the season, are all deciduous. The leaves of Aesculus
calif ornica and Sambucus glauca are the highest in silica-free ash, and
especially in calcium at maturity, of the species in this group. Moreover,
the foliage of A. calif ornica increases greatly in sugars toward maturity,
reaching 8 per cent when the leaves have dried, a factor which may ac-
count for its relatively high palatability late in the autumn. Quercus
Douglasii has the lowest nutritional rank of the four species of this
group. In silica-free ash this plant is considerably the lowest of the four
species. The percentage of silica in these species increases markedly with
advancement of the season, amounting in the mature leafage to nearly
one-half of the ash content, a phenomenon which apparently accounts for
the harshness of the foliage, and the fact that its palatability declines in
the later growth stages. Moreover, the phosphorus, calcium, and potas-
sium contents in Q. Douglasii are lower, whereas the fiber content is
somewhat higher, than in the other three deciduous species. Rhus diver-
siloba is fairly high in nutrients throughout the season, and compares fa-
vorably in this respect with A. calif ornica and 8. glauca. All four species
are of importance as browse because of their fairly high level of protein
and essential inorganic constituents late in the season when the herbace-
ous cover generally is at its lowest nutritional ebb.

COMPARATIVE LEVELS OF CONSTITUENTS
IN THE SIX PLANT GROUPS

Although the trends in composition of the samples of individual species
have been shown, it is of interest to know whether the forage of all the
species embraced in each of the various distinctive plant groups exhibits
characteristic compositional trends and levels. For purpose of compari-
son three widely different growth stages were selected, namely, the early
leaf developmental stage, the flowering stage, and the period of full
maturity of the leafage. Graphic data have been prepared from averages
of all the species of each of the six plant groups.

In deriving the points on the curves an average value for a given
growth stage for each year was first obtained ; then these curves for the
three years were in turn averaged for each species ; and the figures so
derived were then averaged for each plant group. In the half -shrubs and
the nondeciduous shrubs, however, because of the nature of their growth,
the graph points were obtained by computing the averages for each spe-
cies on a calendar basis for the three years, the month of March repre-
senting the initial point because of the abundance of early leafage at that

76 University of California — Experiment Station

time. Data on odd samples, such as flowers and seed, were not included
in deriving these averages.

Figure 19 shows that the deciduous shrubs and trees are much the
highest of the groups in the percentage of crude protein at the early leaf
stage, and also at the time of flowering. Moreover, a relatively high pro-
tein level is maintained in this group at maturity. The broad-leaved herbs
rank second in protein content in the early leaf stage and in the blossom-
ing period, but the grasses and grasslike species show only slightly lower
levels than the broad-leaved herbs throughout the growth cycle. The half-
shrubs are high in protein content in March, followed by a sharp decline,
which reaches the minimum level late in the summer, after which there
is a continuous rise, giving the curve the appearance of a circular seg-
ment. The nondeciduous species maintain a more constant level of pro-
tein throughout the growth cycle in the mixed foliage, being the lowest of
all the groups early in the season, but second highest late in the autumn.

In crude fiber (fig. 20) , the half -shrubs are the highest from late spring
until late summer, when, because of a larger proportion of leafage on the
shoots, the fiber content decreases. The grasses and the grasslike species,
which have nearly the same values, outrank all the other groups in this
constituent, except the half-shrubs, for a part of the season, as pointed
out. Next to the grasses, the grasslike plants, the broad-leaved herbs, the
deciduous shrubs and trees, and the nondeciduous shrubs, respectively,
rank in the fiber levels in the order named. The range in values in these
groups is Large. The nondeciduous shrubs and the deciduous shrubs and
trees run parallel in this constituent throughout the season; they vary
little in values, and are conspicuously lower than the other groups.

The silica-free ash (fig. 21), is highest in the broad-leaved herbs of
the six groups until midsummer, when the deciduous shrubs and trees
take the load. The grasses and the grasslike species maintain nearly the
same level in ash content, but they are distinctly lower than the broad-
leaved herbs. The deciduous shrubs and trees are relatively low in ash
in the early leaf stage ; but the rise continues gradually throughout the
season, reaching the highest point of all the groups late in the autumn.
The half-shrubs and the nondeciduous shrubs form irregular curves of
ash values, the crests and depressions of which are evidently determined
by the proportion of new and old leaves contained in the samples. The
highest crest of the ash level in the half-shrubs comes in the late summer,
whereas that in the nondeciduous shrubs is recorded in early spring.

The calcium content varies widely in the different plant groups. The
broad-leaved herbs lead in this constituent in the early leaf stage (fig. 22) .
Fairly early in the season, however, the deciduous shrubs and trees, fol-

Bul. 627]

Composition of Foothill Plants

77

-

1 1

-

Ok

ZEGEWD

\ GRASSES

*V, _ _ GRASSLIKE PLANTS

*V BROAD-LEAVED HERBS

*V^ DECIDUOUS SHRUBS &• TREES

*S - HALF-SHRUBS

"

_ "S*SS NONDECIDUOUS SHRUBS

■

\-. -% 'Bk

-

<Xs%o^ ^-- v\

■

N^^C"^ \

-

-

\ ^^^v^ •*>. *v,

-

-

-

* * X.... ^0\ \ /

-

-

\^ ^>^v. >-,., "

-

-

^~----^^^S>^. \

-

"~~"~~-^^\^

-

"^-^v\^..

-

-

^^o^.

-

~

^

-

-

i i

-

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 19. — March of crude protein in the six plant groups.

1 ' o

.

___——-- — " — ~^Z-""°

.

-

rr-~~—~~ — "Z^^"

-

-

#'"' — ^^^^'^ s*

"

-

/ ^^^■' '" '" ^°^~—. s'

-

Jb~~ «^, 'L^^^ •■"'" ^>: °

"

-

-

—

/ ^^^'^^■•"' ■'

-

-

L^~ — — **""" -^

.

o — '' »■»■ — S

-

■T

-

-

"

■

—

—

-

J „ ■"

.

-

©

-

-

"

-

-

©-•■

"

-

Q_^

-

-

....-•° G"--.... ..© o .'" """"" *~°

-

-

....©••' ©••••-" © G

—

.

<*•""" LEGEJVD

.

r.DASSF.<!

GRASSLIKE PLANTS

.

. BROAD-LEAVED HERBS

_

.

DECIDUOUS SHRUBS & TREES

.

HALF-SHRUBS

-

NONDECIDUOUS SHRUBS

-

-

1 1

-

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 20. — March of crude fiber in the six plant groups.

78 University of California — Experiment Station

' '

"

"•-•.^

""•"♦.^

"

"""-•s,.

la

_

"""-.^

—

"

•>». ©

"

-

•Sp. __--"'

"

-

"*^- —~ — "

-

10

-

"**••„. "~

-

"

^■"'~X*--~^

"

©..„ _.-•- «"***" *"-■-..

.

a

CX^>^^r,-;:-*-'^"' ^""^

-

^-""""""'^^"^-Cl"'***"--

"

\ ^^^-^^"^-^ ..©

"

\ ..• •■©^^^^^■^•^.^ / "'"■©•-..

"

£

o....... • K ''^^^

-

\ " .^^ /

-

4

^^ ^^><1^

-

LEGEND ""~ o "*'" ^~^~^^

-

f.CAS<!F5 ^

-

GRASSLIKE PLANTS

BROAD-LEAVED HERBS

DECIDUOUS SHRUBS & TREES

"

HALF-SHRUBS

-

"

NONDEC1DUOUS SHRUBS

"

O

1 1

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 21. — March of silica-free ash in the six plant groups.

GRASSES

GRASSLIKE PLANTS

BROAD-LEAVED HERBS

CIDUOUS SHRUBS & TREES

HALF- SHRUBS

:ciduous SHRUBS

/
/■'
/'

S

s

/'
/'

...0'"' ,y

**"■©•-. . .---*''
— o

©

J L

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 22.— March of calcium in the six plant groups.

Bul. 627]

Composition of Foothill Plants

79

lowed by the nondeciduous shrubs, exceed the broad-leaved herbs in this
constituent. At maturity the deciduous shrubs and trees far outrank all
other groups in the percentage of calcium. The grasslike species are the
lowest in this constituent, followed closely by the grasses. The half-
shrubs and the nondeciduous shrubs form irregular curves, reaching the
maximum calcium values in the autumn. The nondeciduous shrubs rank
second in calcium levels from early summer until the following growth
period.

In phosphorus the range in the early leaf stage is clearly the widest of
the three ash constituents studied, and it is the narrowest at leaf matu-

TABLE 10

Trends of the Various Constituents in the Plant Groups,

from the Early Leaf Stage to Maturity

Plant groups

Ash

Calcium

Phosphorus

Potassium

Protein

Fiber

Grasses

Decrease

Decrease

Decrease

Decrease

Decrease

Increase

Grasslike plants

Decrease

Level varies
slightly

Decrease

Level varies
slightly

Decrease

Increase

Broad-leaved herbs . . .

Decrease

Level varies
slightly

Decrease

Decrease

Decrease

Increase

Deciduous half-

Decrease

Level varies
slightly

Decrease

Decrease

Decrease

Nondeciduous shrubs

Level varies

Level varies

Level varies

Level varies

Level varies

Level varies

slightly

slightly

slightly

slightly

slightly

slightly

Deciduous trees and

shrubs

Increase

Increase

Decrease

Decrease

Decrease

slightly

rity, as shown in figure 23. The deciduous shrubs and trees are much the
highest in phosphorus content in the early leaf stage, and the nondecidu-
ous shrubs and the half -shrubs are the lowest throughout the season. In
the broad-leaved herbs, the grasses, and the grasslike groups, the phos-
phorus values are nearly parallel in their decline throughout the season,
the first-named being the highest and the grasslike species the lowest.

In potassium content (fig. 24) the broad-leaved herbs rank first, fol-
lowed by the grasslike species, the grasses, the deciduous shrubs and
trees, the half -shrubs, and the nondeciduous species, in the order named.
The range in potassium content in the various plant groups is wide
throughout the season, and conspicuously so early in the growth period.

The comparative study of the six plant groups brings out several points
of contrast, namely: (1) the levels of the various constituents differ
widely throughout the season in the respective plant groups ; (2) in pro-
tein and phosphorus the range in the levels is highest in the early leaf

80

0.9

University of California — Experiment Station

ae-

go.5

1 ■■■ 1

.

-

V LEGEND

\ ORASSES

-

-

\ . GRASSLIKE PLANTS

-

_

N-N _.. BROAD-LEAVED HERBS

_

\ . DECIDUOUS SHRUBS & TREES

N-N __ HALF-SHRUBS

\ NONDECIDUOUS SHRUBS

-

-

-

"--•-.

-

' U^L "* **

\ "^''^^^rr^ii--.

-

\ -a-^""""""***"^;^

-

z^*8^ "^^^

-

rf °-.\ ^>-^s§

vo.... ,--".-— •••••.;-o

X ©•••.... jr -..-- '--o

^"^ ""-a.. ..S™

^~ e-jri-'^

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 23. — March of phosphorus in the six plant groups.

! 1

"*>

"

\ LEGEND

"

—

GRASSES
"V GRASSLIKE PLANTS

—

^ BROAD-LEAVED HERBS

SS^.. '*% DECIDUOUS SHRUBS & TREES

^\^--.^ V. HALF-SHRUBS

^\^>-.^ v- .% NONDECIDUOUS SHRUBS

"

^^\ "*' ~" - ****"•

"

-

^V. " "- ^

-

^\ "" -"

"

^\ '"^^

^^ "*iv»»,

-

-

«v ^^^ ^ ^.. •

-

"~^^ "^s^N^ ""-^

"

\ " ^ ^ ""^^^-^

*

'

" ^ -^^^\

""^-^ ^^^

...-* -K '^..

-

® "" o""::-^--------^^:-:-::' o- ...

-

"■o

-

-

-

1 1

-

EARLY LEAF FULL BLOOM LEAFAGE DRY

Fig. 24. — March of potassium in the six plant groups.

Bul. 627] Composition of Foothill Plants 81

stage and lowest at leaf maturity; (3) in calcium and fiber the range is
widest at leaf maturity; (4) the leafage of the deciduous shrubs and
trees is relatively high among the groups in protein, phosphorus, and
calcium, and is relatively low in crude fiber and potassium ; (5) the non-
deciduous shrubs are comparatively low in all constituents, except cal-
cium; (6) the half -shrubs are low in protein, silica-free ash and its
constituents, and are high in fiber; and (7) the broad-leaved herbs are
approximately five times higher in calcium than the grasses and grass-
like groups, slightly higher in protein, phosphorus, and potassium, and
much lower in fiber.

A resume of the march of the constituents in the six plant groups dur-
ing the growing season is given in table 10.

APPLICATION TO RANGE MANAGEMENT

Since these studies show that species and groups of species, exhibit char-
acteristic chemical compositions, and that these change continuously and
markedly with their seasonal development, the question arises as to the
significance of this knowledge to practical range problems. Accordingly,
the following inquiries are pertinent :

At what growth stage may the greatest energy and nutritive values
of the different plant associations be procured, without jeopardy to the
range ?

What role of economic importance, if any, do the unpalatable or
"weedy" species play in a range plant community ?

Should the management of a range be so directed as to restore and
maintain as nearly as possible the original plant cover, linked, as it is,
with its characteristic soil type, or may there be a less planned utilization
of the forage, resulting in the vegetation being naturally transformed
into a single plant association, such as a purely herbaceous stand, or into
a brush cover ?

In what ways may knowledge of the march and the levels of the chemi-
cal composition of forage species serve in the recognition of the char-
acteristics especially desirable in a forage species ?

Is it possible to predict the effect of a shortened growing season, such
as may be induced by drought or unfavorable temperature, on certain
specific nutritional values of the forage crop in such years?

May tables giving the chemical values of the dominant species at im-
portant growth stages, and the time involved in reaching these periods, be
used, in conjunction with the survey of the botanical composition of the
range, with a view to refining the estimates of its carrying capacity, as
when a grazing reconnaissance is made ?

82 University of California — Experiment Station

The discussion which follows, taken up in the order of questions raised,
aims to point out how the results reported in this bulletin may throw some
light on these and similar questions.

Comparative Forage Values of the Different Growth Stages. — Since
these studies have shown that the composition of the range species
changes greatly from the early leaf stage to maturity, it seems pertinent
to compare, through the adoption of some standard, the values of the
more essential constituents in the field samples at definite growth stages.
For purposes of comparison 25 pounds14 of oven-dry weight of forage
was taken as a standard weight in each of the three growth stages of the
forage, namely, the early leaf stage, the flowering stage, and at full ma-
turity. Table 11 gives, in pounds, the crude protein, crude fiber, silica-
free ash, and phosphorus content in field samples equivalent to 25 pounds
on the dry-weight basis as above stated. The object of the following
computations is to show the respective nutritive values of forage at
various growth stages when the water factor has been eliminated. Such
comparisons should clarify the question of relative values of the young
and the more mature forage.

Table 11 shows that in the early leaf stage the weight of the field sam-
ple equivalent to 25 pounds of dry material is much the highest, and the
weight of the mature sample is much the lowest, whereas in the bloom
period the weight is intermediate. In the grasses and broad-leaved herbs
it will be noted that in the early leaf stage, despite the fact that t he equiv-
alent to 25 pounds of oven-dry weight of the field sample is much the
largest — owing to high water content — the nutritional values are the
highest for the entire season. Even when the blossom stage lias been
reached, and the moisture content has declined considerably, the daily
poundage of the more essential constituents consumed by the animal,
except in such annual grasses as Festuca megalura and Arena barbata,
is still satisfactory. At maturity, on the other hand, when the field sam-
ples approach closely the weight of the oven-dry unit, the percentage
composition of nutrients is the lowest, whereas in the period of innores
cence these values for most of the species are intermediate.

The deciduous species listed in table 11 show thai the poundage of
crude protein, computed on a 25 pound dry-weight ratio, is highest in
the early leaf stage, and lowest, late in the autumn. At the latter period
the poundage of protein amounts to about one-fourth of that contained
in the young foliage. On the other hand, in the nondeciduous species, as
typified in Ceanothus cuneatus, the protein ratios change little through

14 Twenty-five pounds of dry forage was taken for convenience in these computa-
tions, and also because this figure approaches the daily requirement of a 1,000-pound
animal.

Bul. 627]

Composition of Foothill Plants

83

the season. In crude fiber the ratios in both of these plant groups show
little variation through the grazing period, and is not a determining
factor in their seasonal values.

The silica-free ash in the foliage of the deciduous species is greatest
when the leaves are mature, whereas in the nondeciduous species, the ash

TABLE 11

Proportion of Constituents in Kepresentative Field Forage Samples at

Different Growth Stages, when Converted to the Equivalent

of 25 Pounds of Oven-dry Weight*

Species

Growth stage

Water,
per cent

Approx-
imate
forage

equiva-
lent,

pounds

Silica-
free
ash,

pounds

Phos-
phorus,
pounds

Crude
protein,
pounds

Crude

fiber,

pounds

Avena barbata.

Bromus mollis .

Festuca megalura.

Melica imperfecta .

Juncus oxymeris .

Er odium botrys.

Ceanothus cuneatus .

Aesculus calif or mica.

Quercus Douglasii.

Sambucus glauca.

Early leaf

Flowering

Mature, dry

Early leaf

Flowering

Mature, dry

Early leaf

Flowering

Mature, dry

Early leaf

Flowering

Mature, dry

Early leaf

Flowering

Mature, dry

Early leaf

Flowering

Mature, dry

Early leaf (Feb.)

Fully developed (May)
Leaf hardened (Nov.)..

Early leaf

Flowering

Mature, dry

Early leaf

June

Mature

Early leaf

Flowering

Mature, dry

77.0

71.0

5.0

83.0
75.0
0.3

73.0

62.0

2.3

76.0
75.0
29.0

83.0

71.0

3.0

87.0

86.0

4.0

54.0
54.0
46.0

80.0
78.0
9.0

72.0
61.0
37.0

83.0

83.0

8.0

109
86
26

147
100
25

93
66
26

104
100
35

147

86
26

192

178
26

54
54

125
114

27

40

147
147

27

2.0
1.1
1.3

3.0
1.4
0.7

1.7
0.8
0.4

2.2
1.8
0.8

2.1
1.5
1.3

3.5
2.9
2.6

1.4
1.3
1.4

1.7
2.5

1.6

2.3

2.6
3.8

0.09
.09
.06

.12
.11
.04

.10
.07
.03

.10
.10
.03

.11

.06
.02

.10
.12
.05

.04
.04
.03

.18
.12
.06

.10
.06
.04

.22

.15

0.02

5.3

1.8
0.5

5.6
3.0
0.9

3.8
1.9
0.7

6.0
5.2
0.8

6.0
2.1
0.9

3.2

2.8
2.7

7.6
5.8
1.7

4.9
3.5
2.4

9.1
9.6
1.7

4.7

6.2
7.5

10.6

6.3
7.4
7.9

5.6
8.5
9.0

2.7
4.2
7.1

2.4
3.1
2.6

2.4
2.3
2.9

4.4
3.6
3.2

2.4
2.8
2.4

* Data from samples of different years were used in some cases to obtain the figures for comparative
growth stages.

84 University of California — Experiment Station

fraction is greatest in early spring-. In both groups, however, the pound-
age of phosphorus in a unit of dry weight is highest in the young leafage
and lowest at leaf maturity.

It is significant that when the grasses and the other herbaceous species
are near or in the flowering stage, the crude protein, ash, and the phos-
phorus contents are still fairly high. Despite the fact that the forage is
richest in most constituents in the very early leaf stage, grazing exten-
sively, or to a point which would approach the full capacity of the range
at this period, is for other reasons inadvisable. Very early seasonal graz-
ing, especially if the photosynthetic surface is cropped down repeatedly,
tends to decrease the yield of the season's forage crop. Repeated early
grazing curtails not only the top growth bnl also the rool development.
To obtain the best general results on a herbaceous range the time to bo-
gin grazing appears to 1"- between the earlier growth sta-jv and the flow-
ering period, when the roots have become well established and are ao1

readily Irani pled niit of a wet or loose Boil.

E6U of Weeds in a Rangt Community. Range species, because of
the rigorous environmenl to which they have long been subjected, have
I mat id it about various adaptive responses, which have resulted in fitting
them into their respective ecological niches. The struggle between the
many plant species in a given range area is keen indeed, and results in
the elimination of the major proportion of the initial seedling stand. In
averaged counts of a Bquare fool of surface, which i^ believed t«» be fairly
typical of a well-stocked herbaceous cover of the San Joaquin and similar
foothill range, more than 3,000 Beedlings, composed of many species.
were recorded early in the Beason. < Obviously, not all individuals of such
a dense young stand <-<>nld Burvive. Only the better-adapted forms could
complete their life cycle.

Among the factors which determine the survival of Individuals, the
soil sol in inn appears to play a major part. Preliminary study of the soils
i>\' various plant communities of the San Joaquin Experimental Range
revealed thai the soil solution is generally of \*'vy low concentration. Ac-
cordingly, those species which have the highesl absorptive capacity may
be the ones which determine the ultimate character of the stand.

Prom analytical data presented, it is seen that species which grow side
by side differ greatly in the percentage of the various elements. Tims in
Amsinchia D tal silica-free ash content in the early

growth stages is approximately 20 per cent of the oven-dry weighl of
til-1 plant ; and the potassium content approximates 7 per cent figures
which are conspicuously higher than those in other species invest igated.
Accordingly, thisspe< much greater absorptive capacity than most

Bul. 627] Composition of Foothill Plants 85

i
of its associates. Being largely unpalatable to livestock, it adds much
organic matter and essential salts to the soil, factors which should reflect
favorably upon the establishment and growth of associated species of
quite different absorptive capacities. From the viewpoint of favorably
influencing the soil for incoming plant species, then, some of the so-called
"weedy" species may have a bearing in shaping the character of the plant
community.

Soils in Relation to Plant Types and Forage Utilization. — Most range
areas support a variety of forage species, but often these are composed
largely of a single life form, such as grass or brush. The data here re-
ported show that each of the various plant groups is characterized by
distinctive nutritional patterns. The groups differ not only in the vari-
ations of the constituents throughout the growing season, but also in
their levels.

"Whether the soils, which support these different types of vegetation,
are chiefly responsible for the distribution and the nutritional character-
istics of the various plant groups on a given range is a subject for exten-
sive investigation. The authors, however, have digressed from the main
study to make limited examination of the soils in the sampling sites. It is
hoped that the mere recognition of this phase may stimulate serious work
on this complex problem.

From table 12 it is seen that the soil of a mixed cover of shrubs, trees,
and herbs is richer in total nitrogen and in organic matter than is the soil
of the pure grass stand. Areas sparsely vegetated with Lessingia germa-
norum and Festuca megalura, both of which are shallow-rooted (and
inferior as forage) , were found to have the lowest total nitrogen, nitrates,
and organic material of the soil types investigated. The soil of the wet
swale areas, on the other hand, with its dense and luxuriant growth of
succulent grasslike species, while very low in nitrates, compares favor-
ably with that of the shrub-tree-herb community in total nitrogen and
organic matter ; this indicates low nitrifying power. In these swales, how-
ever, the total concentration of the soil solution was found to be higher in
the deeper soil horizon, whereas the total organic matter was the greater
in the upper soil layer, where the roots are largely found. The pH values
of the soils worked with are only slightly on the acid side, except in the
shrub-tree-herb community, where they drop to 5.99.

It is worthy of note that the soil samples from the wet swale are char-
acterized by a highly selective botanical composition, only a few species
composing the bulk of the cover. Along the margin, where the drainage
is good, the cover is composed of a combination of annual and perennial
herbaceous species. During the three years of study these border species

86

University of California — Experiment Station

have not been observed to invade the true swale type to any appreciable
extent. Since the soil of this area contains only a trace of nitrates (table
12), and is saturated with water during much of the growing season, the
aeration is poor — factors evidently of sufficient magnitude to prevent
invasions by species of the adjoining dry-land vegetation.

In the moist, but well-drained meadow, which is conspicuously char-
acterized late in the season by the robust Hemitonia virgata, and a

TABLE 12

Eelation of Some Soil Characteristics to Plant Cover

ox the San Joaquin Experimental Range

Cover where sample was taki'n

Depth of
soil sample,
incl

Loss on
ignition,
per oenl

nitrogen,

per rent

Nitrate
nitrogen,

p. p.m.

pH

Mixed cover of trees, brush, grass,

f 0-8
f 0-8

( 0-8

i n
i u

in

. 103

100

.053

3.0
0.7

(i 5

trace

0.5

i o

0.5
4 0

6 30

Swale of graaelib

Grass cover with scattered, bi

6.62
6.80

6.57

■pane oover

6.50
6.64

0 ii

scattering of TrifoUum tridentatum and i rptu setigerus, there

are to be Pound occasional robusl Bpecimensof the species which dominate
the marginal and the well-drained adjoining slopes. Since this meadow
soil is well aerated, nitrification is presumably active, and there is ample
\< » . amounting to some 1 parts per million. Moreover, the total organic
mailer and total salts are of much tlic same magnitude as in the s«>il of
the shrub-tree-herb association. Evidently, therefore, it' it were nol for
the kern competition Be1 ap by // la virgata, and the few other

dated species, the adjoining hillside vegetation would presumably
completely occupy this an

These two examples of closed, or BO-called "unyielding" communities,
illustrate well Borne of the more eonspicuou * which determine the
delimitations of plant associations. In the wel swale of the grasslike cover,
the poor aeration ami the general character of the soil evidently deter-
mine the plant population. End 1. even if this cover were removed the

boring hillsi imably could not come in. These swale

ten, serve as indi t distinctive soil characteristics. The

Bul. 627] Composition of Foothill Plants 87

Hemizonia virgata area, on the other hand, illustrates the play of quite
a different set of factors in the determination of the stand. Here the
character of the soil is probably much less a potent factor than is the pres-
ence of the exceedingly aggressive and well adapted dominant species
which compose the present vegetation. If, in this site, the more distinctly
meadow type of vegetation were removed, it is reasonable to assume that
the adjoining hillside species would occupy the area.

In the light of these plant behaviors it appears that the soil solution,
in combination with other edaphic factors, notably moisture supply, may
have an important bearing on the ultimate character of the plant cover.
The subject of sequence of species on developing soils, or on areas where
the upper soil horizon is being gradually removed, appears worthy of
critical study.

Criteria for Judging the Value of a Forage Plant. — Knowledge of the
march of the chemical composition of range plants makes possible the
segregation of the species into two broad categories : those having rela-
tively high nutritive values ; and those of consistently low values. Al-
though such information is valuable, it does not afford a sufficiently broad
index to segregate the species into their respective economic categories.
Aside from the factor of palatability, some species which are high in the
essential constituents may be of relatively little value because of their
short growth cycle. Species like Lessingia germanorum, because of the
low levels of their nutrients throughout the growth period, can be classed
as undesirable purely on the basis of chemical studies. Festuca megalura,
on the other hand, when in the early leaf stage, ranks in nutrients with
those of desirable grass species ; but the life cycle of this species is so
short as to place it among the most undesirable of range grasses. Thus
the criteria for judging the economic rank of a palatable range plant is
that of high nutritive value maintained through the life cycle, prefer-
ably in addition to a long period of vegetative growth. Most of the peren-
nial grasses investigated mature relatively late in the season, a behavior
commonly associated with high nutritiveness well into the autumn.
Knowledge of the seasonal change of composition of the forage species,
therefore, must be considered in addition to data pertaining to the
length of the growth cycle of the individual species, in classifying them
as to relative range values.

Shortening of Growing Season as Affecting Nutritive Values. — It is
not uncommon, particularly in the foothill and valley regions of the
state, that deficient autumn or winter rains, or unfavorable temperature,
may retard the inception of growth by several weeks. Since the present
study, although based solely on nursery plots, appears to show that a

88 University of California — Experiment Station

short growing season results in lower values for phosphorus, and in some
species, for calcium, and in somewhat higher levels for crude fiber, it
would appear that such abnormal years would be associated with poorer
quality of forage.

Although the present investigation has not included vegetation of the
high mountain ranges, the results here reported lead one to suspect thai
the forage there produced, associated as it is with a short growing season,
might be low in phosphorus. Especially may this be the case in years
when the snowfall is so heavy as appreciably to delay the beginning of
growth.

Chemical Values and Grazing Capacity. — The present reconnaissance
methods used in estimating the grazing capacity of a range admittedly
are in need of refinement. Failure to obtain uniformly reliable est imates
may partly be accounted for by the fact that the nutritive value of the
forage changes from month to month and dees not reach some degree of
stability until late in the summer. Also the data obtained for carrying-
capacity estimates do not take into account the rclat ive quanl it ies of the
constituents of the different forage types, and therefore do not take into
account the nutritional balance in the forage as a whole.

Possibly -razing capacity estimates could be refined by taking into
account the following factors: botanical composition of a forage type,
expressed in weighl of air-dry herbage ; the average palatability of each
plant group, such as grasses, broad-leaved herbs, browse, etc.; the aver-
age Length of t lie growing season of each forage type ; and the averaged
march of composition in each planl group. Tims it' it is assumed that one
acre of ;i grass type, when in the early growth stage, produces one-fourth
ton of air-dry material. 90 percent of which is palatable, then 450 pounds
of such forage could be nt Ilized. The protein content, as read from a pre-
pared nutritional curve, is taken ;is 20 per eenl for purposes <>f calcula-
tion, which -jives 90 pounds of this constituent for the area. Estimate- of
weighl increase in the air-dry forage ^ the season advances, accom-
panied as ii is by ;i corresponding decline in crude protein, could readily
!"• calculated from the curve for the type at any time in the season. In

the same way the nutrient values for ;i unit area COUld be derived for

other important constituents. Repetition of this method for each type
would furnish data as to the total nutrients available at any time in the
season. With the inclusion of such data in the reconnaissance computa-
tions would appear to furnish more exacting estimates of the carrying
capacity of ;i range. Tins deducl ion is offered with the hope that a quali-
fied reconnaissance worker may become sufficiently interested to include
in his equation the factors above enumerated.

Bul. 627] Composition of Foothill Plants 89

SUMMARY

The study of foothill range plants, with special reference to the seasonal
march in their composition, was conducted on field and nursery sample
plots of the San Joaquin Experimental Range. The dominant vegetation
is believed to be fairly characteristic of the extensive savanna-woodland
association.

Review of the literature here cited reveals the fact that much confusion
exists relative to the changes in composition of forage species. The fact
that there is a high percentage of crude protein, and a low percentage of
crude fiber in the early growth stages, followed by gradual decline in the
former and increase in the latter with advance in the season, has already
been fairly well established; but the trend in the ash content, and its
constituents, during the life history of range plants, requires further
investigation. Some significant data on this phase are reported in this
paper.

The samples of most species were collected at distinct growth stages,
irrespective of chronological dates; the majority of previous workers
have collected the samples according to established dates.

In most of the graminaceous, grasslike, and broad-leaved herbaceous
species there is a continuous and rather orderly decline in the crude pro-
tein, the silica-free ash, calcium, phosphorus, and potassium from the
earliest appearance of leaf blades to plant maturity. The percentage of
crude fiber in these plant groups, on the other hand, increases with the
advance of the season. The most rapid changes in constituents take place
from the early leaf development to the period of full bloom.

Although species within a plant group have characteristic composi-
tional levels, the variation in chemical values in the entire life cycle of
plants of a given species is greater than that between the various species
of the plant group for any single growth stage.

The levels of protein in the three herbaceous groups (grasses, grasslike
plants, and broad-leaved herbs) at corresponding growth stages are of
similar values ; but the levels in crude fiber in the broad-leaved herbs are
lower, on the average, than in the two other herbaceous groups.

In silica-free ash, calcium, and potassium, the broad-leaved herbs
maintain a higher level throughout the entire season than do the grasses
and grasslike species. In phosphorus content the values in these three
groups are much the same at corresponding growth stages. The grasslike
species are characterized by high silica content.

The crude protein in the foliage of the deciduous shrubs and trees de-
clines in an orderly way from the earliest appearance of the leaves to

90 University of California — Experiment Station

their maturity. The protein levels in this group are the highest of the
various groups studied. In the deciduous half -shrubs and the nondecidu-
ous shrubs the protein levels change relatively little, and are the lowest
of the six groups. The crests and depressions of the curves of these two
groups are determined by the relative amounts of the young and mature
foliage in such vegetation.

The crude fiber in the foliage of the deciduous shrubs and trees, and in
the nondeciduous shrubs, maintains nearly the same level throughout the
season, and is the lowest in any of the groups studied. In the deciduous
half -shrubs the level of the fiber while variable, is the highest of these
three groups, the maximum amount being reached in midsummer.

In silica-free ash, and in calcium content, the foliage of the deciduous
shrubs and trees stands out in contrast to all the oilier plant groups in
that the trend is distinctly upward with advancement in the season, and
these constituents arc much higher than in the other two groups of woody
plants. In the aondeciduous shrubs and the deciduous half-shrubs, the
ash and calcium contents vary throughout the season, the levels being
governed by the proportion of young foliage in the sample. These two
constituents are the Lowest in the deciduous half-shrubs.

The trend in potassium contenl in the three groups of woody plains is
distinctly downward with increasing age of the foliage. In contrast to
the herbaceous species, the potassium contenl lakes second place quanti-
tatively to that of calcium.

The phosphorus contenl declines gradually from the early leaf stage
to leaf maturity. The phosphorus levels of the deciduous and nondecidu-
ous half-shrubs are the lowest of any of the plant groups studied.

There is an impressive consistency in the composition of all constitu-
ents within a species through the growth cycle from year to year.

me of the "weedy," or Largely onpalatable species, have a high ab-
sorption capacity and are conspicuously rich in certain essentia] ele-
ments. Accordingly some such species may play an important part in

determininir the character of the plant cover in subsequenl years.

The calcium-phosphorUS ratio shows characteristic behavior in the

respective plant groups. In th< es this ratio remains close to 1 to 1

at all -row ih stages; in the grasslike species it is about L to 1 in the early

leal' Stage and 4 to 1 at maturity ; in t he broad deaved herbs the average

ratio is approximately J to 1 early in the season and 12 to 1 at late ma-
turity ; in the nondeciduous shrubs the rat io is approximately 5 to 1 in
the early leaf Btage and 16 to 1 late in the autumn. In the deciduous

shrubs and trees the ratio vanes most, averaging approximately 1 to 1

in the early leal Btage and 2fi to 1 at leal' maturity.

Bul. 627] Composition of Foothill Plants 91

Shortening of the growing season apparently results in a decrease in
phosphorus, and in some species in calcium, but it does not influence
quantitatively the protein and potassium contents. Enrichment of the
soils concerned, as through application of manure, on the other hand,
results in a higher percentage of crude protein and phosphorus, and a
lower percentage of crude fiber.

The seeds of the grasses and the broad-leaved herbs, as compared with
the mature leafage, have approximately the same values in calcium,
whereas the phosphorus content is higher, and the potassium content is
lower. The seeds of the shrubs and trees, on the other hand, are lower in
calcium than the mature foliage, whereas the levels of the phosphorus
and potassium are nearly the same as in the foliage.

These studies reveal that a valuable range plant, among other things,
must have a long vegetative period, or the nutritional values of the ma-
ture herbage must be maintained at a high level well after seed forma-
tion. This viewpoint is helpful in the classification of range plants as to
their economic values.

For a given unit of dry matter the early season growth has a higher
caloric value, and is richer in ash and its constituents, than the mature
leafage.

Knowledge of chemical values of range species, especially if considered
in relation to their respective soils, should prove helpful in explaining
plant succession.

A diversified plant cover is preferable for stabilization of nutritive
forage values to that of a single plant type.

Knowledge of the chemical values of the dominant species, with their
seasonal changes should prove useful as a factor in perfecting more de-
pendable methods for estimating the carrying capacity of the range.

ACKNOWLEDGMENTS

The writers wish to make grateful acknowledgment to the following per-
sons, all of the California Forest and Range Experiment Station of the
United States Forest Service: To Station Director E. I. Kotok, and to
M. W. Talbot, Senior Plant Ecologist, for valuable suggestions in the
organization of the work plan and in making available laboratory and
other facilities at the San Joaquin Experimental Range ; to Dr. Harold
H. Biswell, Assistant Conservationist, for invaluable assistance in col-
lecting forest samples and for help in many other ways ; and to Assistant
Station Director Dr. J. A. Hall for critically reading the manuscript.
The spirit of cooperation and helpfulness is greatly appreciated.

92 University of California — Experiment Station

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