CHARACTERISTlCS OF SCOTCH PINE PROVENANCES
RELATED TO EUROPEAN PINE SAWFLY ATTACK

Thesis for the Degree of M. S.
MICHiaAN STATE umvsnsm’
Wiiliam K. Randail
19.65

 

mags LIBRARY

Michigan State
University

 

 

am ESE ow

William K. Randall

CHARACTERISTICS OF SCOTCH PINE PROVENANCES

RELATED TO EUROPEAN PINE SAWFLY ATTACK

ABSTRACT
In 1958 seeds were requested from native Scotch pine (Pinus

_§y1vestris L.) stands in several parts of Europe and Asia. Response

 

was good and seedlots were obtained from 108 native stands plus a
number of plantations. In 1961 the NC-51 series of test plantations
was established. From 40 to 108 different seedlots were planted in
each plantation; the planting following a randomized complete block
design. In the years 1963 to 1965 four test plantations in southern

Michigan suffered attack by the European pine sawfly (Neodiprion

 

sertifer (Geoffrey)).

This study is part of a long-range Scotch pine improvement
project. The objectives were: determine genetic differences in
sawfly resistance in Scotch pine, and determine the factors responsible
for possible resistance.

The damage estimates made in the years 1963 to 1965 consisted
of counts of the number of infested trees. From each tree three
1-year-old needles were collected and measured in July 1965. The
collections were made from centralbrlocated branches on which feeding
had occurred.

Slow growth was probably responsible for the limited attacks on

the most northern origins -- the insects could not find the trees in

William K. Randall
the weeds. However, it is doubtful whether growth rate was a major
factor in resistance among the medium and fast-growing varieties. Also,
there were instances in which a tall tree of var. uralensis was free of
damage whereas a nearby smaller tree was heavily eaten.

Several factors were possibly correlated with sawfly resistance.
Early onset of growth, wide needles, slow to medium growth rate,
intense autumn coloration, and early needle maturity were characteristics
of those seedlots with the fewest attacks. Those seedlots were mostly
from northern latitudes. They also had 11mmu' concentration of foliar

sodium, potassium, and magnesium than did susceptible provenances.

CHARACTERISTICS OF SCOTCH PINE PROVENANCES

RELATED TO'EUROPEAN PINE SAWFLY ATTACK

by

William K. Randall

A THESIS

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

MASTER OF SCIENCE

Department of Forestry

1965

ACKNOWLEDGEMENTS

My sincere gratitude is given to Dr. J. W. Wright (Forestry
Department, Michigan State University) who devoted considerable time
advising and reviewing the procedures of this study. Thanks are
given to Mr. Walter Lemmien, and those of the W. K. Kellogg Forest,
who kindly assisted in the collection of data. Many thanks to
Louis Wilson, Klaus Steinbeck, and John Ruby, whose findings were
incorporated within this study. Finally I wish to thank those who
gave of their time in review and suggestions throughout the course

of this study.

ii

TABLE OF CONTENTS

INTRODUCTION . . . . . . .
PAST WORK IN BREEDING FOR INSECT-RESISTANCE

Resistance and cause .
Examples of resistance .
Permanence of resistance .

MATERIALS AND METHODS

Kellogg Forest

Russ Forest
Allegan Forest

Rose Lake .
Needle measurements

SCOTCH PINE

Distribution .

Description

Importance . . . . . . . . .
Evolution within the species

EUROPEAN PINE SAWFLY .

Occurrence .
Description
Life history .
Damage .
Oviposition

RESULTS

Date of onset of growth

Needle hardness

Needle width .

Needle color . . .

Tree height . . . . . . . . . . .
Other possible resistance factors

iii

Page

12
12
13
13

16

16
16
21
21

24

24
25
25
25
26

28

28
28
31
32
32
38

TABLE OF CONTENTS (continued)
Page
DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

Future tests are needed . . . . . . . . . . . . . . . . . . 46
Future breeding program . . . . . . . . . . . . . . . . . . 46

SUMMARY . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

APPENDIX . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

iv

Table

LIST OF TABLES
Page
Varieties (and numbers of origins) of Scotch

pine (Pinus sylvestris L.) included in the test
and their natural ranges (from Ruby, 1964). . . . . . . . 22

 

Date of onset of Scotch pine growth and the
percent of mature needles per tree, related to
European pine sawfly resistance . . . . . . . . . . . . . 29

Scotch pine tree height and foliage character-
istics possibly related to resistance to the
European pine sawfly . . . . . . . . . . . . . . . . . . . 30

Relation between sawfly attack and height of
Scotch pine varieties in southern Michigan,
1965 . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Scotch pine foliar potassium, sodium, and
magnesium as related to attack by the European
pine sawfly . . . . . . . . . . . . . . . . . . . . . . . 44

LIST OF FIGURES
Figure Page

1. Locations of the outplantings (solid dots)
which were sampled for sawfly attack . . . . . . . . . . 11

2. Natural distribution of Scotch pine in Europe
(shaded) and provenances included in Wright
and Bull (1963) test (numbered dots). . . . . . . . . . . 18

3. Natural distribution of Scotch pine in Asia
(shaded) and provenances included in Wright
and Bull (1963) test (numbered dots) . . . . . . . . . . 20

4. a.--Pinus sylvestris var. mongolica
(Yakutskaya, Siberia 60° 45' N. Lat. 131°
40' E. Long.) 6 years from seed, Rose Lake
Wildlife Experiment Station, Shiawassee
County, Michigan. Trees this small were
rarely attacked . . . . . . . . . . . . . . . . . . . 35

 

b.--Pinus sylvestris var. haguenensis (Moselle,
France 49° 36' N. Lat. 2° 06' E. Long.) 6
years from seed, Rose Lake Wildlife
Experiment Station, Shiawassee County,
Michigan. This and other seedlots of the
same variety were heavily attacked by the
sawfly . . . . . . . . . . . . . . . . . . . . . . . 35

 

 

5. Pinus_§ylvestris var. hercynica (Bohemia,
Czechoslovakia 50° 12' N. Lat. 15° 3' E. Long.).
A susceptible provenance defoliated by the
European pine sawfly. Photo taken at the Rose
Lake Wildlife Experiment Station, Shiawassee
County, Michigan . . . . . . . . . . . . . . . . . . . . 37

 

6. Pinus sylvestris var. uralensis (Ural
Mountains of Russia 56° 51' N. Lat. 610 23’ E.
Long.). A resistant provenance planted at the
Rose Lake Wildlife Experiment Station,
Shiawassee County, Michigan . . . . . . . . . . . . . . . 4O

 

7. Relationship between height and trees attacked
for the northern, central, and southern groups
of Scotch pine varieties . . . . . . . . . . . . . . . . 42

vi

INTRODUCTION

No farmer today would think of planting an orchard of wild apples,
or a field of the inferior forms of corn from which the present very
productive hybrid strains were developed. Increasingly, the forester is
aware of the possibilities of improving forest trees and is trying to
replace unimproved wild types with more productive strains.

There is genetic variation in all species. Certain individual
trees are able to grow better on a particular site than are others.
They grow faster than their neighbors and leave more seed. On another
site other types of trees grow faster and produce more seed. Thus, in
the course of time, natural selection results in the development of
different races.

The racial variation pattern of Scotch pine (Pinus sylvestris L.)

 

is well known through a series of studies going back to 1820. Most of
the data pertains to growth characteristics. Some of the racial

studies have been conducted in southern Michigan. Recently, some of the
Michigan test plantations were infested by the European pine sawfly

(Neodiprion sertifer (Geoffrey)). This provided an opportunity for the

 

study of genetic differences in resistance to attack by this important
pest.

My study is part of a long-range Scotch pine improvement project.
My objectives were two: determine genetic differences in sawfly
resistance in Scotch pine; and, determine the factors responsible for

possible resistance. To this a third objective may be added as part of

the long-range project -- breed a resistant strain with satisfactory
growth characters.

There are five commonly recognized methods of achieving insect
control. They are:

1. Direct control by spraying with insecticides.

2. Biological control by the introduction of parasites or

predators.

3. Ecological control by making the environment unfavorable for

the insect.

4. Radiation control by the sterilization of males which are re-

leased in large numbers and cause females to lay sterile eggs.

5. Genetic control through the breeding of resistant tree

varieties.

Insecticides have been sprayed over millions of acres as an insect
control practice. Life is contaminated when the invisible insecticidal
shroud settles on the earth's surface. Biological, ecological, and
genetic control are methods for reducing insect populations without
producing undesirable side effects which may accompany direct control.
In the future we must always remember that our forest resource is not
composed of timber alone and that other values must also be preserved.

A sound knowledge of the factors causing immunity or resistance and
the conditions which influence these factors, is the basis for breeding
plants for insect-resistance. However, it is not necessary to under-
stand thoroughly these factors in order to make progress in a breeding
program.

Genetic control through breeding has obvious advantages and

disadvantages. In the case of this particular pest, it may cost many

thousands of dollars to produce a resistant strain of Scotch pine. But
once produced, the strain would cost no more to plant than an ordinary
one and would eliminate the need for periodic spraying of thousands of
acres. Thus in the long run, genetic control might well be the most

economical.

PAST WORK IN BREEDING FOR INSECT-RESISTANCE

Resistance and cause.--Painter (1951) defined insect-resistance as:

 

"The relative amount of heritable qualities possessed by the plant which
influence the ultimate degree of damage done by the insect." Sdegaard
(1964) defined resistance more simply: "Resistance refers to trees that
are less damaged or less infested than others under comparable
environmental conditions." Resistance results from the presence of a
substance or structure in resistant plants and its absence in suscepti-
ble ones, or the reverse.

A knowledge of the cause of resistance is highly desirable, but it
may or may not be of use in breeding programs. The agronomist does not
demand a full knowledge of the cause of a high yield before breeding
for this character in field crops. It is no more necessary to know the
exact cause in breeding for insect resistance. In the majority of the
breeding programs for insect resistance, the exact cause of resistance
has remained unknown (Painter, 1951).

Breeding for forest tree insect-resistance is relatively new. Most
of the literature cited for this study is by R. H. Painter, E. J.
Schreiner, and various workers from the Institute of Forest Genetics,
Placerville, California. Painter’s interest is breeding for insect-
resistance in agricultural plants. His methods, however, are applicable
to breeding for insect-resistance in forest trees. Painter's
recommended procedures were as follows:

1. A survey for possible sources of insect-resistant varieties

and strains.

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2. .A determination of some of the basic properties of the plants
responsible for resistance.

3. Hybridization to combine genes for resistance with desirable
agronomic characters.

4. A study of genetics of resistance to the insect where possible.

5. Testing the resistance in advanced-generation hybrids.

6. The study of resistance of released varieties in plots and on
farms to evaluate resistance as an insect-control method.

Examples of resistance.--Cherry fruit flies (Rhagoletis cingulata

 

 

Loew) are unable to lay their eggs in cherries that have a certain de-

gree of hardness (Snelling, 1941). This may be influenced by either

the stage of maturity of the fruit or varietal characteristics.
Soft-wooded Chrysanthemums have been reported to be more suscepti-

ble to Paroxuna micella Lw. than the hard-wooded varieties. Painter

 

(1951) found that fiber hardness is a character which is associated
with resistance in certainxarieties of sugar cane to the sugar cane

borer (Diatraea saccharalis Fab.).

 

A stiff-strawed Pawnee variety of wheat (Triticum aestivum L.)

 

has been developed which is resistant to the hessian fly (Phytopaga

destructor Say.). European grape (Vitis vinifera L.) is susceptible to

 

 

the grape phylloxera (Phylloxera vitifoliae Fitch) while American

 

varieties are resistant (Painter,1951).
R. C. Hall (Forest Insect Laboratory, Columbus, Ohio) from 1930

to 1940 found two clones of black locust (Robinia pseudoacacia L.)

 

resistant to the locust borer (Magacyllene robiniae Forster): (l) the

 

resistant 'Shipmast' clone found on Long Island, New York; (2) the

'Higbee' clone of southern Indiana.

 

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Schreiner (1949) conducted an experiment breeding poplar. He found

differences in resistance to the Japanese beetle (Popillia japonica

 

Newman). Four interspecific hybrid combinations demonstrated that there
are wide differences between siblings of the same hybrid combination.
The combinations in which this differential resistance occurred were:
Populus 'charkowiensis' X P. balsamifera var. virginiana Foug.
'charkowiensis' XIP. 'caudina'

'charkowiensis’ X.P. berolinensis Dipp.
. simonii Carr. X'P. berolinensis Dipp.

 

 

 

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In each case one parent was itself an interspecific hybrid.
The Institute of Forest Genetics, Placerville, California, has de-
veloped an insect-resistant variety of pine, by crossing Jeffrey pine

(Pinus jeffreyi Grev. and Balf.) with Coulter pine (P. coulteri D. Don.)

 

then using pollen from the hybrid and backcrossing to Jeffrey pine. The

backcross hybrid was resistant to the resin midge (Retinodiplosis spp.)

 

and the pine reproduction weevil (Cylindrocopturus eatoni Buch.). In

 

resistant pines the cortical tissue became necrotic around the egg
puncture made by the female pine reproduction weevil. In trees which
were killed the larvae penetrated the necrotic tissue and reached the
cambium. The larvae are unable to penetrate the necrotic tissue in
resistant trees. Miller (1950) stated that resistance was possibly
attributable either to this necrotic layer or to some property of the
resin.

Yellow-green varieties of chick peas (Cicer arietinum L.) are more

 

resistant to the pea aphid (Macrosiphum pisi Harris) than blue-green

 

varieties (Painter, 1951). The European larch (Larix decidua Mill.)

 

is susceptible to the larch gall aphid (Chermes spp.) which is not

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1964).)

Permanence of resistance.--Insect-resistant varieties impose ad-

 

verse effects on the insect's life cycle when used as host. These
resistant characters act as control mechanisms in reducing the potential
insect population, thus resulting in lesser host damage. However, with
a resistant variety the factors of permanence and degree of resistance
must be considered. Permanence of resistance depends upon several
factors (Painter, 1941).
l. The proportion of acreage of a resistant to susceptible
varieties in a given area.
2. The thoroughness with which other control measures are
practiced.
3. The purity of the resistant variety as grown.
4. The number of genetic factors and resistance characters
involved in the resistant variety.
5. Genetic relationships of the strain of insect to other
insects.
6. The original proportion, if any, of the population capable of
feeding on the resistant variety.
7. Ecological adaptation of the strain of insect feeding on the
resistant variety.
Ninety years ago phylloxera-resistant grape vines were sent from
the United States to France. These vines still are an important means
of control of the grape phylloxera in Europe. Ordinary cotton

(Gossypium spp.) could not be grown in South Africa because of the

cotton leafhopper (Empoasca fascialis Jac.). However, a hairy variety

 

of (Gossypium hirsutum L.) has been found which is not attacked. This

 

hairy variety of cotton has remained since 1925 (Painter, 1958).

Varieties of wheat (Triticum aestivum L.) resistant to the wheat

 

rust (Puccinia graminis Pers.) have been developed (Shaw, 1964).

 

However, new biological strains of the fungus appear and the formerly
resistant wheat variety is rendered susceptible. Perhaps four years is
the total maximum resistant period for any new wheat variety. This
unfortunate situation results because the diploid stage of the life

cycle occurs on the barberry (Berberis canadensis Mill.) while the

 

haploid stage is confined to the wheat plant. The haploid stage is
capable of rapid genetic change. In a diploid organism considerable
more time is needed to produce a true-breeding new type. Most insects
are diploid. Hence we do not expect the rapid evolution of damaging

new types as has occurred in the wheat rusts.

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MATERIALS AND METHODS

In 1958 seeds were requested by J. W. Wright from native Scotch
pine stands in several parts of Europe and Asia. Response was good and
seedlots were obtained from 108 native stands plus a number of phanta-
tions. The seeds were sown in the former Michigan State University
Bogue Research Nursery in 1959 and the seedlings were grown there for
two years and under the direction of W. 1. Bull.

In 1961 the NC-51 series of test plantations was established.
There were 10 such plantations in various parts of Michigan plus a
number in other north central states. From 40 to 108 different seedlots
were planted in each plantation; the planting followed a randomized
complete block design. The number of replications varied from 7 to 10.
An 8-by—8 foot spacing and 4-tree linear plots were used in each case.

In the ensuing years four of these test plantations suffered attack
by the European pine sawfly and could be used in this study. These four
were located in southwestern Michigan (Figure 1). More complete
descriptions of these follow.

Kellogg Forest.--Plantation 2-61 is located on the W. K. Kellogg

 

Forest, Kalamazoo County, 2 miles west of Augusta, Michigan. There are
108 Scotch pine origins represented in 10 replicates totalling about
4800 test trees. Planting strips were furrowed in October to remove the
sod. Seedlings were hand planted in the furrows the following spring
under the direction of W. L. Lemmien.

Two replicates are on level areas, the others are hilly with slopes

to 40 percent. The soil is moderately fertile (Oshtemo loamy sand). The

10

Figure 1.--Locations of the outplantings in southern Michigan
(solid dots) which were sampled for sawfly attack.

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12
level areas were cropped prior to planting. Survival as of 1964 was 93
percent. Average plantation height at that time was 34 inches. During
the 1965 growing season many of the trees grew an additional 20 inches.

Near the test area is a 30-year-old Scotch and red pine (Pinus resinosa

 

Ait.) stand, the probable source of sawfly infestation.

Russ Forest.--Plantation 7-61 is located on the Fred Russ Forest,

 

Cass County near Dowagiac, Michigan, and was planted under the direction
of J. W. Wright and John Bright. It contains 105 Scotch pine origins
replicated 10 times, a total of 4435 experimental and border trees. Chemi-
cal weed control was used to eliminate competing vegetation in the planting
strips. Survival in 1965 was 83 percent. The area is level and the sandy
loam is fertile. The plantation area was used as a nursery prior to plant-
ing. Average plantation height in 1964 was 34 inches.

Light sawfly infestations in 1963 and 1964 probably came from

planted ponderosa (Pinus ponderosa Laws.) or Scotch pine a quarter mile

 

distant. Nearby areas were sprayed by airplane for another insect pest
in 1965. That apparently destroyed all sawflies.

Allegan Forest.--Plantation 11-61 is located on the Allegan State

 

Forest, Allegan County near Allegan, Michigan. It was planted under the
direction of J. W. Wright. There are 72 origins in 10 replications for a
total of about 3000 experimental trees. It was machine planted with 2-0
seedlings. Scalping was unnecessary because of the slight amount of
ground cover which consisted largely of lichens, poverty grass (Danthonia

spicata (L.) Beauv.) and prickly pear cactus (Opuntia humifusa Raf.).

 

The planting site is level. Soil is coarse textured and relatively

infertile. Survival as of 1965 was 88 percent. Average plantation

13

height in 1965 was 37 inches. Red pines (Pinus resinosa Ait.) 30 feet

 

tall, a probable source of sawfly infestation, are located about 200
feet east of the test plantation.

Rose Lake.--Plantation 12-61 is located on the Rose Lake Wildlife
Experiment Station, Shiawassee County near Lansing, Michigan. It con-
tains 75 origins and 7 replications, or about 2100 test trees. It was
established by machine with no weed control. John L. Ruby and J. W.
Wright did the planting.

The plantation site is rolling with up to 10 percent slopes., The
soil is well drained loamy sand with a sparse cover of lichens, poverty
- grass and dewberry (Rubus sp.). A clay layer at'a depth of 2 to 3 feet
‘provided good moisture relations once the trees were established. In
1964 the average heightwas 34 inches with some origins as tall as 45
inches. The taller origins grew an additional 2 feet in 1965. Survival
as of 1964 was 90 percent.

There are two east-west rows of Scotch pine about 30 feet south of
the test plantation. These trees average 15 feet in height. These
trees have been severely attacked by the sawfly and undoubtedly provided
the source of infestation for the test plantation.

The test trees suffered a moderate amount of sawfly damage in 1963.
All infested test trees were sprayed so that new attacks would have to
come from the neighboring rows of trees. This same practice was
followed in 1964, when the attacks were heavier. The 1965 infestation
was very severe.

Needle measurements.--I used previously gathered data on the per-

 

centage of first-year trees having mature needles and summer foliage

14

color of 2—year—old seedlings. In both instances the information was
gathered from the nursery test. The data on foliar analysis were
supplied by Klaus Steinbeck, based on samples collected at the Russ
Forest plantation in the autumn of 1963.

The damage estimates made in 1963 and 1964 by Louis F. Wilson and
J. W. Wright consisted of counts of the number of infested trees. Tree
counts were also made in 1965. In addition the amount of damage per
tree was determined. At Kellogg and Allegan the number of sawfly
colonies was counted on each tree. At these two plantations the percent-
age of foliage eaten was rarely more than 5 percent. At Rose Lake the
amount of damage was determined by estimating, to the nearest 5 percent,
the amount of foliage eaten.

Needles for measurement were chosen from a branch on which feeding
had occurred. Three l—year—old needles were collected from each tree
in July 1965. Each needle was from a different fascicle. The
collections were made from centrally located branches on which feeding
had occurred. In this way it was hoped that the measurements would
apply to needles as the insect saw them.

The needle measurements were made in the laboratory with a dissect-
ing microscope. The lengths were measured to the nearest millimeter
and the widths to the nearest 40 microns.

Leaf hardness was estimated with the aid of a dissecting needle
pressed against the epidermis by hand. It was hoped that this would
duplicate the action of an ovipositor. Five grades of hardness were
recognized. Possible observer bias was eliminated by hiding the

identity of the leaf until the observation was recorded.

 

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Each set of measurements for a single test plantation was subjected
to analysis of variance. The degrees of freedom were as follows: S-l
for seedlot, R-l for replicate, and (S-l)(R—l) for error, S and R being
the numbers for seedlots and replicates in a plantation respectively.

No transformations were used.

 

 

SCOTCH PINE

Distribution.--Scotch pine occurs naturally over an extensive area

 

of Europe and Asia (Figures 2 and 3). The most northerly occurrence is
near Alten, Norway (70° N. latitude). From there it ranges southward

to the Sierra Nevada Mountains (37° N. latitude) of the Andalusia region
of southern Spain. The Sierra Nevadas also mark the western boundary
(50° W. longitude) of its range. The eastern extreme is delineated by
the Aldan River (137° E. longitude) of northeastern Siberia. In its
native range Scotch pine is one of the most important commercial timber
species. It grows on varied sites, from the cold-climate

pinetum-cladinosum cover type of northern Eurasia to the warmer mountain

 

types of southern Europe. It is by preference a tree of siliceous soils,
but occurs on almost all geological formations (Wilde, 1958).

Description.--Wright and Bull (1963) portrayed Scotch pine thus:

 

"Needles 2 per cluster 1 to 3-1/2 inches (25 - 80 mm.) long, 1/25 to
1/12 inch (1-2 mm.) broad, stiff, sharp-pointed, twisted, gray-green,
with persistent needle sheaths; cones 1-1/2 to 2-1/2 inches (4-6 cm.)
long, 2 to 3 times as long as broad, conic with a rounded base, yellow
to grayish brown, with long angular apophyses on the basal scales,
opening in December or January and not long persistent after ripening;
bark scaly, dark brown or black at first but changing to yellow or

reddish on large branches or on the lower boles of large trees."

16

17

Figure 2.--Natural distribution of Scotch pine in Europe (shaded)
and provenances included in Wright and Bull (1963)
test (numbered dots).

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Figure 3. Natural distribution of Scotch pine in Asia (shaded)
and provenances included in Wright and Bull (1963)
test (numbered dots).

 

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,i \
000. 00'. con. bN. b: 08. k b. bk .0.

 

 

 

21

In the Scottish woods the Scotch pine is flat-topped and only 50
to 60 feet tall whereas spire-crowned trees 100 to 120 feet tall are
common in old German and Scandinavian forests.

Fully stocked Scotch pine stands do not have as heavy a canopy as
most other northern coniferous stands. Therefore, profuse ground
vegetation occurs under 30- and 40-year-old stands.

Within any population that is transcontinental in distribution,
there is apt to be a great amount of genetic differentiation. Scotch
pine is transcontinental. Carlisle (1958) named lll geographical and
morphological variants. Those included many cultivars of limited im-
portance, and there was some duplication among names. Ruby (1964) used
genetic and taxonomic methods in arriving at his classification of the
species. He used data from Michigan growth tests as well as measure-
ments taken on cones, seeds, and leaves collected from wild trees. He
recognized 21 geographic varieties (Table 1), each characterized by
differences in growth as well as morphological characteristics.

Importance.--Scotch pine is an important species in the United

 

States. It is the most important exotic planted today in the northeast-
ern and north central regions of the country. Its use, for the most
part, has been limited to Christmas trees. However, in the near future
pulping of this species may become important. This importance is shown
by the numbers of trees grown in Michigan nurseries. As of 1963 there
were 28 million seedlings in private nurseries within the state. About
14 million seedlings are planted in Michigan each year.

Evolution within the species.--During the Pleistocene Epoch vast

 

glaciers scoured the present geographic range of the Scotch pine.

Nerthern Europe and northern Asia were subjected to glaciation which

 

l
x .
J.
..
.
.i
.
- ..
-.
.
1A
A
A A
L.
a \. .—
-

 

J

 

22

Table l. Varieties (and numbers of origins) of Scotch pine (Pinus
sylvestris L. ) included in the test and their natural ranges

 

(from Ruby, 1964).

 

lapponica (5)
mongolica (2)
uralensis (4)
altaica (3)

septentrionalis (16)

rigensis (5)
armena (8)
aquitana (6)
scotica (4)
iberica (5)
nevadensis (0)
rhodopaea (5)
illyrica (1)
pannonica (2)
carpatica (l).

vindelica (0)
engadinensis (O)

borussica (2)
hercynica (20)

scotica X ? (2)

polonica (2)

haguenensis (10)

Northern Norway, Sweden, and northwestern USSR
Eastern Siberia

Ural Mountains in USSR

Altai Mountains in USSR

Central and southern Norway, central Sweden, and
central Finland

Southern Sweden and Baltic countries

Northeastern Turkey, Armena, Georgian SSR, and Iran
Massif Centrale of France

Highlands of Scotland

North-central Spain

Sierra Nevada Mbuntains of southern Spain

Southern Bulgaria, northern Greece

Central Yugoslavia

Western Hungary

Northeastern and eastern Czechoslovakia

Southeast central France, Switzerland, and western
Austria

Engadine Alps, upper Inn Valley, and eastern
Switzerland

Northeast German lowlands
Southern and eastern Germany, central Austria

Plantations in England probably originating as
hybrids between Scottish and south European trees

Poland

Western Germany, eastern France, and Belgium

 

J
1A
,
J.

t.

i

' 1 IL.
I,

J
‘ L
o

A
a.
.

23

eradicated most of the vegetation. However, there were a few isolated
refuges where segments of the populations survived. Wright and Bull
(1963) postulated that five such places existed --"the Ural Mountains
of Russia, the Pyrenees of France and Spain, the Kjolen Mountains of
Norway and Sweden, and the Alps or Carpathians of southeast Europe? In
pre-Pleistocene times the pine was probably well differentiated so that
these remnants formed distinct races. It is believed that the present
varieties descended from these remnants.

When a barrier prevents the mingling of new characters as they
arise, separated populations sooner or later evolve along divergent
lines due to selection pressure. Thus there is an independent accumula-
tion and loss of genes within each separated population. However, in
the continuously forested areas of Germany and Czechoslovakia no
barriers existed. Therefore, little differentiation resulted in such
areas. Changes due to selection pressure are much slower than those
due to environmental changes. Thus Scotch pine in some locations is
not adapted to its modern environment. To a certain extent varieties
of Scotch pine reflect their Pleistocene ancestry (Wright and Bull,

1963).

.ll]l||111

 

 

EUROPEAN PINE SAWFLY

Occurrence.--The European pine sawfly, a Hymenopterous leaf—eating

 

insect, was introduced from Europe. It was first collected in North
America in Somerset County, New Jersey, 1925. The present known range
in the United States encompasses the northern states from New England
to Michigan and southern Ontario. The sawfly also ranges to Iowa,
southern Illinois, and southern Ohio. In Europe it is distributed in
Sweden, Finland, Russia, Germany, Austria, Hungary, Czechoslovakia, and
occurs in Japan.

According to Ross (1955) the original Neodiprion group, which
spread across North America with its conifer host during the Mid-
tertiary or early Miocene (40 to 50 million years ago) was later split
into eastern "lecontei" and western "sertifer" groups by the formation
of the Great Plains. European pine sawfly or its ancestral form later
dispersed to the Eurasian continent via a Bering land bridge.

The European pine sawfly attacks Scotch pine, red pine (P.

resinosa Ait.), Japanese red pine (P. densiflora Sieb. and Zucc.),

 

jack pine CE. banksiana Lamb.), table-mountain pine (P. pungens Lamb.),
and mugo pine (P. mugo Turra.). Also there is slight feeding on eastern
white pine (P. strobus L.), Austrian pine (£-.Ei§£§ var. austriaca
(Hoess) Aschers and Graebn.), ponderosa pine QB. ponderosa Laws.),
shortleaf pine (P. echinata Mill.), and pitch pine (P. rigida Mill.)
(Lyons, 1964). Occasional larval feeding has been reported on black

spruce (Picea mariana (Mill) B.S.P.) and whitespruce (Picea glauca

 

 

(Moench) Voss.) when these trees grow in close proximity to pines.

24

 

 

f..-

25

Although no one host species is consistently preferred by ovipositing
females, Scotch pine is more suitable for the development and survival
of the larvae (Lyons, 1964).

Description.-Full grown larvae are 3/4 to 1 inch (20 to 25 mm.) in

 

length, with a black head and a gray-green body having a gray
longitudinal stripe. The adult male is 1/4 inch (6 mm.) long, black in
color with a feather-like antennae. The adult female is 3/8 inch (9
mm.) long, yellowish-brown in color with a thread-like antennae.

The oval egg is pale yellow-white, 0.05 inch (1.3 mm.) long by 0.01
inch (0.13 mm.) wide. The cocoon is a light tan or dark golden brown,
cylindrical with rounded ends, 0.3 to 0.4 inch (8 to 10 mm.) long. A
male cocoon is slightly smaller than a female cocoon.

Life History.--The European pine sawfly completes its life cycle in

 

one year. Winter is passed in the egg stage on the needles. Hatching
in mid-April or May, the larvae feed for a period of four or six weeks.
In mid-June or July the cocoons are spun in the duff beneath the in-
fested trees. The prepupae remain in their cocoons until August or
early September at which time the adults emerge. After mating the
female deposits her eggs in slits cut in the edge of the current year's
needles.

Griffiths (1959) noted that most pupae emerge as adults in late
summer or early fall; a few overwinter in the cocoon for one or more
years. This survival of overwintering pupae serves to buffer the popu-
lation against the catastrophic elimination of a single class-year.

Damage.--Defoliation causes a reduction of height growth. Wilson

(1964) found that 10 larval colonies per tree feeding on 5-foot-tall

1

L4

 

26

Scotch pines in southern Michigan reduced height growth by 14 percent
during the following year. Under those same conditions 25 larval

colonies per tree reduced height growth by 23 percent. Seldom does a
5-foot-tall tree support more than 25 larval colonies on the average.

Oviposition.--Egg laying occurs in late August or early September.

 

A female flies to a tree and selects a needle, usually above the level
of adjacent weeds. She uses her thread-like antennae to determine the
presence of eggs already laid. Then the female backs down the entire
length of the needle before any egg laying is done. However, Ghent
(1959) stated that females may lay eggs along any portion of the needle.

Visualizing a needle from its flat surface, apex upward, a female
cutting egg pockets on the right edge of the needle extends her left
legs only a short distance across the flat surface. The right legs
must reach the greater distance around the curved surface, to grasp the
same edge. In this position she cuts the egg pocket and deposits a
single egg. The female then advances and the process of cutting an egg
pocket and depositing an egg is repeated until 6 to 8 eggs are laid per
needle. Egg laying ceases when the female's antennae detect the
narrowing of the needle at its tip.

Egg laying proceeds while the legs are fully extended across the
needle. In the case of a wide needle the leg movement is restricted
and closer egg spacing results. Thus there is an inverse relationship
between needle width and the egg spacing. A female may find that an ex-
ceptionally wide needle is difficult to grasp so she abandons it and

seeks a narrower one.

27

An inverted position is adopted by the female in 76 percent of the
cases (Ghent, 1959). This is due to the greater stability offered by
having its weight slung below the needle.

Females of all sizes can oviposit on narrow needles, but only the
largest females are able to utilize wide needles, that is, needles over
1.5 mm. (Lyons, 1964).

The female deposits her eggs on several needles, thus forming an
egg cluster. Griffiths (1959) determined that on Scotch pine an egg
cluster contains 90 eggs.

Many Scotch pine in southern Ontario, during the 1958 growing sea-
son, produced exceptionally wide needles (Ghent, 1959). In areas of

heavy infestation these needles were shunned.

RESULTS

Date of onset of growth.--Table 2 gives the day of year that

 

growth started in 1960 for different origins of Scotch pine. Onset of
growth was determined when the terminal bud opened and elongation was
apparent. The less attacked trees started growth two days earlier than
the heavily attacked origins; the difference was significalt (5 per-
cent level). The two-day spread in the start of growth is a character
that has continued and this difference was apparent in 1965.

Table 2 also shows the percentage of trees with mature needles on
October 1, 1959. The wide difference in percent of needle maturity
cannot be accounted for alone by the two-day spread in the onset of
growth. The difference is apparently due to a timing mechanism that
has developed in certain trees, since after the first year the character
of needle maturity was the same for all trees.

Sawfly attack was less on trees with a high percentage of mature
needles at the end of the first year's growth than trees with a low
percentage of mature needles in the years 1963 to 1965. This character
of needle maturity related to insect attack was merely observed and may
be only incidental to resistance.

Needle hardness.--The mean and range of needle hardness for the

 

resistant and susceptible trees was respectively 5.4, 3 to 12; and 6.1,
4 to 9. The lower number signifies the harder needles (Table 3).
When the resistant and susceptible origins are grown in mixture,

the slight difference in needle hardness may be detected by the female

28

 

29

Table 2. Date of onset of Scotch pine growth and the percent of mature
needles per tree, related to European pine sawfly resistance.

 

 

MSFG No. Started Mature needles
and North growth per tree
country Variety latitude 1960* 1959**
Degrees Day of year Percent

 

Resistant origins

 

 

546 SWE lapponica 60.9 113 87
255 SIB mongolica 52.3 113 84
257 URAL uralensis 56.8 113 59
258 URAL uralensis 58.7 114 80
260 URAL uralensis 57.0 114 43
256 SIB altaica 56.7 113 90
230 FIN septentrionalis 60.5 113 52
273 NOR septentrionalis 59.7 113 84
523 SWE septentrionalis 61.2 113 82
Susceptible origins
207 GER hercynica 49.7 115 16
210 GER hercynica 53.1 115 16
308 CZE hercynica 50.2 115 13
312 CZE hercynica 50.9 115 21
525 GER hercynica 50.3 115 11
211 GER polonica 53.8 115 13
252 GER haguenensis 49.3 115 11
318 BEL haguenensis 51.2 115 10
235 FRA haguenensis 48.2 115 10

 

aBELgium, CZEchoslovakia, FINland, FRAnce, GERmany, NORway, SIBeria,
SWEden, URAL Mountains.

*, ** Differences between groups of origins are significant at the 5 and
1 percent levels,respectively.

.fll

 

 

30

Table 3. Scotch pine tree height and foliage characteristics possibly
related to resistance to the European pine sawfly.

 

 

 

 

 

Needle
MSFG No. Tree August
and height Length Width Hardnessb colox‘3
country Variety 1965 1965 1965 1965 1961
Inches “Mm. Microns Grade Grade
Resistant origins
546 SWEa lapponica 22 40 1530 5 3
255 SIB mongolica 27 50 1580 4 6
257 URAL uralensis 54 58 1600 8 7
258 URAL uralensis 42 52 1650 l 6
260 URAL uralensis 42 63 1750 12 8
256 SIB altaica 31 55 1550 7 7
230 FIN septentrionalis 21 38 1500 5 3
273 NOR septentrionalis 33 47 1640 3 3
523 SWE septentrionalis 36 45 1600 4 5
Susceptible origins
207 GER hercynica 59 67 1480 7 12
210 GER hercynica 58 70 1500 9 12
308 CZE hercynica 58 64 1520 4 12
312 CZE hercynica 55 73 1330 7 12
525 GER hercynica 57 61 1440 0 12
211 GER polonica 54 67 1500 5 12
252 GER haguenensis 56 73 1440 6 12
318 BEL haguenensis 62 79 1580 8 12
235 FRA haguenensis 52 56 1490 9 9

 

aBELgium, CZEchoslovakia, FINland, FRAnce, GERmany, NORway, SIBeria,
SWEden, URAL Mountains.

b 0 hardest, 20 = softest

‘3 o

yellowest, 12 = greenest

 

 

31

sawfly. If hardness differences are detectable she probably selects
the softer needle.

The hardness of plant tissue may be a cause of resistance (Painter,
1951). This relationship is a mechanical prevention to feeding or egg
laying. Painter (1951) further stated that it is during the ovipositing
stage that mechanical obstructions are most likely to be a factor of
resistance.

Ghent (personal letter, 1965) posed the theory that slight dif-
ferences of needle hardness between Scotch pine provenances should prove
no deterrent to the ovipositing female European pine sawfly. The
possibility exists that if a hard, wide needle is encountered by the
female sawfly, she may find it relatively awkward to wield the
chitinized saw while her legs are in an unnatural over-extended position.
Then possibly the combination of a hard, wide needle might prove re-
sistant. Ghent (1960) proposed tree selection for needle width and
hardness as an indirect approach to sawfly resistance.

Results of this study indicate that selection for needle hardness
is not justifiable for the selection of resistant trees.

Needle width.--Significant between-origin differences were found in

 

needle width. As a group the resistant origins had wider needles than
the susceptible origins. The mean widths for the resistant and
susceptible origins were 1.60 mm. and 1.48 mm., respectively (Table 3).
There were two susceptible origins with wide needles (MSFG 211 and
MSFG 318. Two resistant origins (MSFG 546 and MSFG 230) had narrow

needles.

 

 

32

Recent studies (Ghent, 1959 and 1960) have indicated that needles
of 2.0 mm. or more in width are virtually immune to the European pine
sawfly. The female sawfly encountering a wide needle that is also
hard may find it awkward to oviposit. She seeks a soft, narrow needle
on which to deposit her eggs. However, if such a needle is not avail-
able, oviposition may take place on the hard, wide needle. The result
may be fewer eggs per needle and fewer clusters per tree. Thus the
resulting damage from the feeding larvae will be less.

Needle color.--There were parallel trends in late summer foliage

 

color and amount of sawfly attack. Resistant origins had yellower
foliage than susceptible ones (Table 3). The color was scored on
August 10 which slightly precedes the onset of the egg laying period.
At that there was a moderately wide range of needle color.

Although color sensitivity of the sawfly is not known, some
insects respond positively to a segment of the color spectrum (Painter,
1951). Peak sensitivity seems to be to wavelengths in the ultraviolet
and the violet range. While the blue and blue-green portion causes
less response, the yellow-yellow-red portion causes even less (Painter,
1951).

Tree height.--Resistant origins of Scotch pine are found in

 

northern latitudes. Their growth is generally slower than in origins
from more southerly latitudes. The mean height for the resistant group
in 1965 was 34.2 inches and the mean for the susceptible group was 56.8
inches (Table 4,Figures 4 and 5).

Provenances MSFG 229 (var. lapponica) and MSFG 245 (var.
mongolica) are resistant to the European pine sawfly, but their growth

is extremely slow. They should not be included in a breeding program

33

Table 4. Relation between sawfly attack and height of Scotch pine
varieties in southern Michigan, 1965.

 

 

 

 

 

Country Trees attacked at

of Rose Mean

Variety origin Kellogg .Allegan Lake Mean Height

- - - Percent of trees - - - - Inches
lapponica SWE FINa 0 o 0 0 19
mongolica SIB 0 0 0 0 21
uralensis URAL 4 1 1 2 43
altaica SIB 5 2 0 2 32
septentrionalis FIN SWE NOR 3 3 12 6 32
rigensis SWE LAT 7 4 38 16 41
armena GEO TUR 6 6 36 16 39
aquitana FHA 11 16 38 22 40
scotica SCO 14 -- 25 -- 36
iberica SPA 20 22 46 29 38
rhodopaea GRE 7 25 29 30 41
illyrica YUG 25 26 56 36 48
pannonica HUN - 25 17 69 37 50
carpatica CZE 10 -- -- -- 54
borussica GER 38 19 67 41 53
hercynica GER CZE 33 22 70 42 53
scotica X ? ENG 26 -- 72 -- 56
polonica POL 28 28 73 43 51
haguenensis GER FRA BEL 36 27 79 47 63
Average 19 15 45 26 48

 

aBELgium, CZEchoslovakia, ENGland, FINland, FRAnce, GEOrgian SSR, GERmany,
GREece, HUNgary, LATvian SSR, NORway, POLand, SCOtland, SIBeria, SPAin,
SWEden, TURkey, YUGoslavia, URAL Mountains.

34

Figure 4a.—-Pinus sylvestris var. mongolica (Yakutskaya, Siberia
60° 45' N. Lat. 131° 40’ E. Long.) 6 years from seed,
Rose Lake Wildlife Experiment Station, Shiawassee
County, Michigan. Trees this small were rarely at-
tacked.

 

Figure 4b.--Pinus sylvestris var. haguenensis (Moselle, France
49° 36' N. Lat. 2° 06' E. Long.) 6 years from seed,
Rose Lake Wildlife Experiment Station, Shiawassee
County, Michigan. This and other seedlots of the
same variety were heavily attacked by the sawfly.

 

 

35

 

36

Figure 5. Pinus sylvestris var. hercynica (Bohemia, Czechoslovakia
50° 12' N. Lat. 15° 3' E. Long.). A susceptible
provenance defoliated by the European pine sawfly.

Photo taken at the Rose Lake Wildlife Experiment Station,
Shiawassee County, Michigan. Age: 6 years from seed.

 

 

37

 

38

for a desirable insect-resistant strain. Rudolf stated (1960):
"Superior trees should be free, or nearly so, of damage from insects or
disease." "Superior" also refers to growth rate and other desirable
tree characters. A slow growing variety even though insect resistant
is not compatible with the program of selection of a superior variety
for the future.

Slow growth was probably responsible for the limited attacks on the
most northern origins -- the insects could not find the trees in the
weeds. However, it is doubtful whether growth rate was a major factor
in resistance among the medium and fast-growing varieties. To test

this one need only compare the large differences in susceptibility be-

 

 

tween the equally fast growing varieties uralensis, regensis, armena,
and rhodopaea (Table 4). Variety uralensis and var. iberica are very
similar in growth rate, however, only 2 percent of the trees of var.
uralensis were attacked while 29 percent of the trees of var. iberica
were damaged. .Also, there were instances in which a tall tree of var.
uralensis was free of damage whereas a nearby smaller tree was heavily
eaten. Thus var. uralensis was best in the respect that it alone
exhibited unique resistant qualities (Figures 6 and 7).

Other possible resistance factors.--The susceptible origins have

 

longer needles than the resiStant ones. The mean for the susceptible
group was 68 mm. and the mean for the resistant group was 50 mm.
(Table 3).

There were significant differences between the resistant origins

and the susceptible ones in their relative amount of sodium, potassium,

39

Figure 6.——Pinus sylvestris var. uralensis (Ural Mountains of
Russia 56° 51’ N. Lat. 61° 23' E. Long.). A
resistant provenance planted at the Rose Lake Wild-

life Experiment Station, Shiawassee County, Michigan.
Age: 6 years from seed.

 

 

40

,x

U
' "I 424".

u ' . l K

. . M.’

i n I 1. .

. .. 1...! ,- x. ,. 4 . 1 >9

. . ~‘ .' . - ,.~. I]; ‘ _‘ ‘

x 7' 'é‘ . I [,— / o‘ «..' ’ 1 «33%. ."..‘l .
.. 7...\~

3‘s " fir " ‘
.|’~. ‘9‘

I
...J .,

(I " .2' - ‘3“ ' 1

.' " h’v' ' . ' '- If; 12‘ :1."
{3.33}! - [4 v'it'.V.:f-?- WI . ‘ ..

I

 

41

Figure 7.--Re1ationship between height and trees attacked for the
northern, central, and southern groups of Scotch pine

 

 

varieties.

Northern varieties Central varieties
lapponica polonica
mongolica borussica

altaica hercynica
septentrionalis haguenensis
rigensis scotica X ?
uralensis pannonica

illyrica
West and South varieties

scotica
iberica
aquitana
rhodopaea
armena

(Siberian and Ural Mountain seedlots circled N's)

 

 

 

CENTRAL \C

C

      

 

TREES AT TACKED . PERCENT

0 O

O

 

 

 

 

 

 

 

 

7O

7-YEAR HEIGHT. INCHES

 

43

and magnesium. The susceptible origins had higher amounts of sodium,

potassium, and magnesium (Table 5).

44

Table 5. Scotch pine foliar potassium, sodium, and magnesium as related
to attack by the European pine sawfly. .

 

 

 

 

MSFG No.
and
country Variety Potassium** Sodium* Magnesium*
Percent Ppm Percent
Resistant origins
546 SWEa lapponica .48 32 .10
255 SIB mongolica .46 48 .06
257 URAL uralensis .54 29 .07
258 URAL uralensis .52 69 .08
260 URAL uralensis .48 32 .07
256 SIB altaica .52 56 .07
273 NOR septentrionalis .52 52 .07
523 SWE septentrionalis .52 56 .07
Susceptible origins

207 GER hercynica .56 60 .08
210 GER hercynica .50 134 .08
308 CZE hercynica .56 64 .10
525 GER hercynica .56 48 .09
211 GER polonica .52 79 .13
252 GER haguenensis .58 60 .10
318 BEL haguenensis .56 56 .09
235 FRA haguenensis .58 93 .07

 

aBELgium, CZEchoslovakia, FRAnce, GERmany, NORway, SIBeria, SWEden,
URAL Mountains

*, ** Differences between resistant and susceptible groups are
significant at the 5 and 1 percent levels, respectively.

I; .
1 J_.
7'1-
l . .2 A
A. ( L 1.

1

. 1
.
.

A

l

l
't

,l

A

.,2 ;..

 

 

 

DISCUSSION

The resistant factors determined in this screening procedure are
related to the adult sawfly. The adult sawfly flies to new areas and
selects the individual trees in which to deposit her eggs. The odor, or
other characters of a tree which attract the adults for oviposition, does
not always correspond with the suitability of the attractive plant for
larval food or survival.

A tree may possess a component of resistance called antibiosis, and
may affect the biology of the insect (Painter, 1958). Lyons (1964)
stated that although no one host species is consistently preferred by
the female European pine sawfly, Scotch pine is more suitable for the
development and survival of the larvae. Insect development on varieties
of Scotch pine was determined by Wilson (Personal communication, 1966).

L. F. Wilson compared European pine sawfly larvae on a resistant origin

 

var. uralensis with a susceptible origin var. haguenensis. All larvae
found on the resistant varieties were one feeding instar behind those
larvae found on the resistant varieties.

The response of insects to chemical attractions and repellents
in plants generally is thought to constitute a major factor in resist-
ance to insect attack. Insects apparently orient to the odorous sub-
stance (Smith, 1960). This is apparent in the hybrids ponderosa X
Jeffrey pine and Jeffrey X Coulter pine. Each produces resins which are

toxic to the western pine beetle (Dendroctonous brevicomis Le Conte) but

 

are non-toxic to the mountain pine beetle (2. monticolae Hopkins).

 

45

 

 

more
pots
to .
pot
Du}
am(
sax
re

UP

t1
01
Si

iv

V3.

at

he
thc
trc

aft

Ira:

46

Future tests are needed.--Future experimentation is needed to learn

 

more about the factors causing resistance. A future test relating
potassium content to insect attack can be conducted by adding potassium
to resistant trees. The addition of potassium would raise content of
potassium in resistant trees to the level of the susceptible ones.
During the following years if there are differences in the relative
amount of attack between the fertilized and the control trees (of the
same origin) then the foliar potassium level is a contributor to sawfly
resistance. Similarly the addition of other chemicals and their effect
upon the European pine sawfly can be tested.

If an occasional single resistant tree of a heavily attacked origin
is found, the needles from that tree can be compared to needles from the
three other trees of the same plot and also to needles rme other trees
of the same origin. If the needles of the resistant tree are
significantly wider than all other needles, needle width may be suggest-
ive as being a resistant character.

Shading prevents the needle color change from green to yellow. If
var. uralensis is shaded and there are differences in the amount of
attack then yellow foliage can be considered a resistant factor.

Tree height can be held constant by controlling or limiting the
height of the taller trees. The faster growing trees can be pruned to
the height of the slower growing trees. An alternate method of con-
trolling height is planting the faster growing trees one or two years
after the slower growing ones were planted.

Future breeding program.--The future breeding program is the

 

transfer of one character (resistance) from one parent and several

 

.l .r'llI
{I’lllillllllllllfl

47

characters (growth and form) from another parent into the genetic
component of the progeny. The Fl's (progeny from the original cross)
are back-crossed to the recurrent parent (the parent contributing
several characters). The Fz's are likewise back-crossed and the program

continues until a desirable sibling is attained.

SUMMARY
Scotch pine varieties lapponica (Norway, Sweden, and northwest
Russia) and mongolica (Siberia) were not attacked by the European pine
sawfly. Varieties uralensis (Ural Mountains of Russia) and altaica
(Altai Mountains of Mongolian Republic) are also resistant. Of these
four resistant varieties, uralensis is the most rapid growing and has
the best form. Incidence of sawfly attack was greatest on variety

haguenensis (Germany, east France, and Belgium).

 

The combination of needle width, needle hardness, foliage color,
number of mature needles, and post-Pleistocene evolution all interacting
are correlated with sawfly resistance. The factors that show possible
resistance are short, mature, hard, wide, yellow-green needles. Con-
versely, those factors possibly related to susceptibility are long,
immature, soft, narrow, blue-green needles. Susceptible origins con-
tained higher concentrations of foliar sodium, potassium, and magnesium.

One inescapable silvicultural requirement is the need to develop
resistant trees that are also well adapted to the climate and the soil
of potential planting sites. A second highly desirable objective is the
development of resistant trees that are also superior in growth and in
timber quality (Austin, 1927; Rudolf, 1956; Meager, 1957). The resist-
ant varieties themselves cannot be called "superior trees." There is
much regular breeding work remaining; crossing resistant X rapid growing
varieties. Perhaps, the result then approaches the superior tree classi—

fication, a desirable, fast growing, and insect-resistant tree.

48

I||

 

49

Future work includes the evaluation of the female sawfly preference
and deterrent factors. Also included in this study should be the
determination of the oleoresin content of the foliage. Are certain
origins resistant only because there is an assortment of origins in all
test plantations? That question is readily answered by caging female
sawflies and determining their egg laying response. Planting the
resistant origins in solid blocks is an alternate experiment. The
latter method,even though time and land consuming is desirable before

recommendations and the release of planting stock are made.

film/Wind}. Era-IE
, ll . .

uIIJl.

REFERENCES

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Forestry 25: 928-953.

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Beal, J. A. 1957. The outlook for selecting and breeding trees
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Cain, S. A. 1944. Foundations of plant geography. Harper and Bros.,
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Carlisle, A. 1958. A guide to the named variants of Scots pine (Pinus
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Clapper, R. B. and J. M. Miller. 1949. Breeding and selecting pest-
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Cram, W. H. and G. E. Brack. 1953. Performance of Scotch pine under
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Dahms, R. G. 1939. Plant breeding and selecting for insect resistance.
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Finlayson, T. 1960. Taxonomy of cocoons and puparia, and their con-
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Gabriel, W. J. 1958. The possibilities of developing strains of white
pine resistant to the white pine weevil. Proc. Soc. Amer.
Foresters. pp. 58-60.

Gatherum, G. E. and K. F. Jensen. 1964. Scotch pine provenance trial
in Southeast Iowa. Iowa State Jour. Sci. 38: 405-413.

Gerhold, H. D. 1959. Seasonal variation of chloroplast pigments and

nutrient elements in the needles of geographic races of Scotch
pine. Silvae Genetics 8: 113-122.

50

 

"ll' I
j’|j’l’
‘(Ill

51

Ghent, A. W. 1959. Row type oviposition in Neodiprion sawflies as
exemplified by the European pine sawfly N. sertifer (Geoff.).
Can. Jour. Zool. 37: 267-281.

 

1960. A study of the group-feeding behavior of larvae of the
jack pine sawfly, Neodiprion pratti banksianae (Roh.). Behavior
26: 110-148.

 

Griffiths, K. J. 1959. Observations on the European pine sawfly
Neodiprion sertifer (Geoff.). Can. Entomol. 91: 501-512.

 

Grimal'skiJ, V. I. 1961. Priéiny UstojEivosti SosnovyH Nasaidenij K
Hvoegryzus'c'im Vrediteljam Zool. ‘z‘. Moskva 40: 1656-1664. [The
cause of the resistance of Scotch pine strains to needle-eating
insect pests.]

Heikkenen, Herman J. 1965. Dendroctonus pseudotsuga: a hypothesis
regarding its primary attractant. Science 150: 1457-1459.

 

Holling, C. S. 1955. The selection by certain small mammals of dead,
parasitized, and healthy prepupae of the European pine sawfly,
Neodiprion sertifer (Geoff.). Can. Jour. Zool. 33: 404-419.

 

Holst, M. and C. Heimburger. 1955. The breeding of hard pine types
resistant to the European pine shoot moth (Rhyacionia buoliana
Schiff.). Forestry Chron. 31: 162-169.

 

Jensen, K. F. and G. E. Gatherum. 1964. Scotch pine provenance trial
in northeast Iowa. Iowa State Jour. Sci. 38: 377-384.

Jordan, D. S. 1906. Concerning variation in animals and plants.
Pop. Sci. Monthly 69: 481-502.

Kriebel, H. B. 1954. Bark thickness as a factor in resistance to white
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Langlet, O. 1959. A cline or not a cline - a question of Scots pine.
Silvae Genetica 8: 13-22.

Leyton, L. 1955.' Mineral composition in the foliage in relation to the
growth of Scots pine. Forest Sci. 1: 210-218.

Lyons, L. A. 1964. The European pine sawfly, Neodiprion sertifer
(Geoff.) (Hymenoptera: Diprionidae). Proc. Entomol. Soc. Ont.
94(1963) (1964). pp. 5-37.

and K. J. Griffiths. 1962. Observations on the development of
Neodiprion sertifer (Geoff.) within the cocoon (Hymenoptera:
Diprionidae). Can. Entomol. 94: 994-1001.

 

 

d. .Id. 517:.leth Q
. . - fl
{Fail .

 

52

Mansford, K. and R. Raper. 1956. The free and combined amino-acids in
some plant juices. Annals Bot. 20(78): 287-295.

McJean, D. N. 1963. Ecology of Scots pine in the Scottish highlands.
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Meagher, G. S. 1957. Some silvicultural aspects of pest control through
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Miller, W. E. and H. J. Heikkenen. 1959. The relative susceptibility
of eight pine species to European pine shoot moth attack in
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Natarajan, A. T-:.E£.El° 1961. Karyotype analysis of Pinus sylvestris.
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Painter, R. H. 1941. The economic value and biologic significance of
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“

 

53

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1957. The 1938 International Union Scotch pine provenance
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1958. Species hybridization in the hard pines, series
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, R. T. Bingham, and K. W. Dorman. 1958. Genetic variation within
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Jour. Forestry 56: 803-808.

 

i
1'

54

and W. J. Gabriel. 1959. Possibilities of breeding weevil-
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and W. 1. Bull. 1963. Geographic variation in Scotch pine.
Silvae Genetica 12: 1-25.

1963. Genetic variation among 140 half-sib Scotch pine
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1964. Hybridization between species and races. Unasylva 18:
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\

Zobel, B. J. 1958. Seed orchards - their concept and management. Jour.
Forestry 56: 815-823.

APPENDIX

55

 

 

Appendix 1. Scotch pine foliar concentrations of nitrogen, potassium,
phosphorus, calcium, magnesium, and sodium sampled at
Russ plantation.
MSFG No.
and
Country Variety N K P Ca Mg Na
------ Percent - - - - - Ppm

546 SWEa lapponica 2.20 .48 .235 .41 10 32
255 SIB mongolica 2.10 .46 .245 .41 .06 48
257 URAL uralensis 1.92 .54 .210 .41 .07 29
258 URAL uralensis 1.88 .52 .210 .32 .08 69
260 URAL uralensis 2.04 .48 .227 .41 .07 32
256 SIB altaica 1.96 .52 .218 .32 .08 29
273 NOR septentrionalis 2.22 .52 .245 .32 .07 52
523 SWE septentrionalis 2.02 .52 .235 .41 .07 56
207 GER hercynica 1.98 .56 .193 .41 .08 60
210 GER hercynica 1.98 .50 .235 .43 .08 134
308 GER hercynica 1.96 .56 .227 .30 .10 64
525 GER hercynica 1.96 .56 .227 .27 .09 48
211 GER polonica 1.84 .52 .193 .46 .13 79
252 GER haguenensis 1.93 .58 .218 .35 .10 60
318 BEL haguenensis 2.06 .56 .235 .30 .09 56
235 FRA haguenensis 1.92 .58 .227 .43 .07 93

 

aBELgium, FINland, FRAnce, GERmany, NORway, SIBeria, SWEden, URAL
Mountains.

57

 

 

Appendix 2. Scotch pine foliar concentrations of manganese, iron,
copper, boron, zinc, aluminum, and molybdenum, samples
taken at Russ plantation.

MSFG’No.

and
Country Variety Mn Fe Cu B Zn A1 Mo
________ ppm _ _ - _ _ _ - _ _

546 SWEa lapponica 1050 98 12.0 33 57 1030 1.8

255 SIB mongolica 1200 102 7.6 31 67 1230 1.9

257 URAL uralensis 1020 85 11.0 31 71 960 1.9

258 URAL uralensis 840 82 9.3 29 50 1200 1.4

260 URAL uralensis 1200 98 7.6 30 60 1320 1.8

256 SIB altaica 1030 121 10.2 34 64 1070 1.4

273 NOR septentrionalis 1200 124 9.3 38 44 980 1.4

523 SWE septentrionalis 1180 121 10.2 39 54 1210 1.9

207 GER hercynica 680 111 9.3 39 83 730 2.1

210 GER hercynica 780 82 8.4 39 60 940 2.2

308 CZE .hercynica 1100 118 8.4 40 44 1090 1.3

525 GER hercynica 1070 98 7.6 29 44 1050 1.2

211 GER polonica 300 66 8.4 23 75 1300 2.2

252 GER haguenensis 1080 91 9.3 41 64 1130 1.6

318 BEL haguenensis 1160 111 9.3 35 47 1070 1.3

235 FRA haguenensis 700 91 8.4 33 64 1010 2.1

 

aBELgium, CZEchoslovakia,

Mountains.

GERmany, NORway, SIBeria, SWEden, URAL

Ii/ ./

/l.|l|. I 1’1 1i II II]- I'll-1

Appendix 3.

Number of Scotch pine trees attacked by the European pine

sawfly in 1963 and 1964.

 

MSFG No.

and
Variety

Kellogg

1963

1964

RUSS

1963 1964

Allegan

1963

1964

Rose Lake
1963 1964

 

lapponica
229
546
547
548
549
Total

mongolica
254
255
Total

uralensis
257
258
259
260
Total

altaica
227
234
256

Total

OIOOOO OIOO OIOOOOO

HIP-”CO

septentrionalis

228
230
232
233
222
521
522
523
524
543
544
545
273
274
201
Total

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59

Appendix 3. (continued) NUmber of Scotch pine trees attacked by the
European pine sawfly in 1963 and 1964.

 

 

MSFG No.
and Kellogg Russ Allegan Rose Lake
variety 1963 1964 1963 1964 1963 1964 1963 1964 Sum
------------ Number - - - - - — - - — - - -
rigensis
542 - 6 - - - - 1 - 7
541 0 3 l 0 0 0 3 4 11
550 l 3 0 0 l 0 - - 5
223 0 2 0 0 0 0 2 2 6
224 _1 _2 .9. .2 .9. __0 .9. .1 .2
Total 2 16 1 0 1 0 6 7 33
armena
213 0 3 0 0 l 0 5 2 11
214’ 0 O 0 0 - - - - 0
220 0 3 0 0 1 0 2 2 8
221 0 3 0 0 0 0 2 2 7
261 0 l 1 0 - - l 1 4
262 0 2 0 1 - - - - 3
263 0 0 0 0 — - - - 0
254 __9 _9 .9. .2 _: _: _: __: _9
Total 0 12 l l 2 0 10 7 33
aquitana »
212 1 4 0 0 - - - - 5
238 2 0 0 0 2 l l 0 6
239 0 3 2 0 3 0 7 4 19
240 O 0 0 0 - - - - 0
249 - - - - 0 0 4 2 6
316 0 0 0 .0 - - - — 0
320 __3. .2. _1 _0 - .; 11 .31. 2.1
Total 6 9 3 0 5 1 23 10 57
scotica
265 0 3 - - - - - - 3
266 0 3 0 0 - - - — 3
267 0 2 0 0 - - l 0 3
253 .3. _f1. _9 .9. _:_ .: __: _: .1.
Total 1 12 0 2 - - 1 0 16
iberica
218 0 7 0 2 0 0 11 4 24
219 0 7 2 2 0 0 3 l 15
245 0 l 0 0 2 l l 1 6
246 l l 0 0 2 0 2 2 8
247 .1 .2 .2 __0 .9 .2 _1. .1. .2
Total 2 20 2 4 4 3 18 9 62

60

 

 

Appendix 3. (continued) Number of Scotch pine trees attacked by the
European pine sawfly in 1963 and 1964.
MSFG-No.
and Kellogg Russ Allegan Rose Lake
variety 1963 1964 1963 1964 1963 1964 1963 1964 Sum
------------ Number - - - - - - - - - - - -
rhodapaea
243 1 1 l 0 l 0 l 1 6
244 0 0 0 1 0 1 6 6 14
271 0 4 0 1 3 4 - - 12
272 0 2 0 1 - - 2 2 7
551 .2 .9 .2 .9 .9 __°. _: .: .9.
Total 1 13 1 3 4 5 9 9 45
illyrica
242 l 6 0 2 2 4 ll 6 32
pannonica
552 3 2 l 2 - - 9 4 21
553 .9 .9 .1 .9 .1 .2 .19 .9. 9.9
Total 3 10 2 7 l 2 24 12 61
carpatica
314 0 l - - - - - - 1
borussica
202 0 6 3 l l l 6 6 24
210 .9 .9 .9 .2. .9 .2 .19. .11 91
Total 0 15 3 3 4 3 20 23 71
hercynica
203 1 4 0 2 l 0 0 2 10
204 0 5 0 l l 1 - - 8
207 l 7 0 4 3 0 9 12 36
208 l 4 0 l 2 1 9 6 24
248 - - - - - - 13 ll 24
525 l 6 l 2 3 0 13 11 37
526 2 8 0 0 - - - - 10
527 0 6 0 1 2 0 ll 5 25
528 2 6 0 0 l 0 - - 9
529 5 12 1 1 - - - - 19
305 1 9 0 0 2 1 ll 18 42
306 l 6 3 l 5 0 9 9 34
307 2 9 0 4 1 3 - - 19
308 2 5 l 4 l 2 8 6 29
309 0 6 0 2 l 0 7 4 20
310 4 7 0 2 1 l 13 13 41
311 0 6 0 2 l 0 10 14 33

L n. .Inn'iquuflwl‘IEJ
I .II'I. .

1" l 1" l

 

61

 

 

Appendix 3. (continued) Number of Scotch pine trees attacked by the
European pine sawfly in 1963 and 1964.
MSFG No.
and Kellogg Russ Allegan Rose Lake
variety 1963 1964 1963 1964 1963 1964 1963 1964 Sum
------------ Number - - - - - - - - - - - -
hercynica (con’t)
312 l 8 2 2 3 0 12 13 41
313 0 l - - - - - - 1
315 l 9 - - - - - — 10
319 .: ..: '2 ..3. .3 .9 9 13 .22
Total 25 124 T6 32 30 9 132 1'3? 501
scotica X ?
269 2 13 0 2 — - 17 15 49
270 .1 _9 ..1_ .3. _-_ .: .: _' .13
Total 3 22 l 5 - - 17 15 63
polonica
211 0 8 0 5 1 1 9 10 34
317 .1. .2. .9 .1. .9 .1. 19 .9 21
Total 1 10 0 6 l 2 19 16 55
haguenensis
206 1 5 1 l 0 2 10 3 23
250 2 8 4 2 - - l6 17 49
251 2 12 0 7 0 0 15 19 55
252 2 3 2 8 l 3 8 12 49
253 4 8 2 3 2 2 14 14 49
235 1 5 2 0 1 1 9 6 25
236 0 8 0 2 0 2 - - 12
237 1 13 8 8 - - - — 30
241 2 10 l 5 0 1 17 15 51
318 0 ll 0 0 0 2 24 19 56
530 .2. 19 .9 .9 .9 .9 .19 .91 .99
Total 17 96 23 39 7 13 131 126 452
no described variety
554 3 3 l 2 — - 6 2 17
555 0 2 3 0 - - - - 5
556 2 1 2 0 - - 7 7 19
557 0 2 1 0 - 7 3 13
225 0 5 0 2 1 0 7 12 27
205 l 4 - - - — - - 5
209 .9. .E .9 .1 .. .: .; _; ._7
Total 6 23 7 5 1 0 27 24 93

62

 

 

 

 

 

Appendix 4. Number of Scotch pine trees attacked by the European pine
sawf1y and the amount of attack per tree, June 1965.
MSFG No. Trees attacked Attack per tree
and Rose Rose
variety Kellogg Allegan Lake Sum Kellogg Allegan Lake
- - - - Number - - - - - - Colonies - - Per-
cent
lapponica
229 0 - - 0 0 - -
546 O O 0 0 0 0 0
547 0 - - 0 O - -
548 0 - - O O - -
549 _2 .: .: .3. .2 ‘ ‘
Total 2 0 0 2 6 "'6 _6
mongolica
254 o o 0‘ o 0 o o
255 .9 .9 .9 .9 .9 0 0
Total 0 0 0 0 0 "6 "6
uralensis
257 4 - 7 11 4 - 15
258 0 1 2 3 O 1 20
259 0 0 3 3 0 0 24
260 .1 ..: .5. .9 .2. - 25
Total 5 l 17 23 6 1 "921
altaica
227 0 0 - 0 O 0 -
234 2 - - 2 4 - -
256 .1 .1. .9 .9 .2. .1 0
Total 3 1 0 4 6 1 —6
septentrionalis
228 0 - - O 0 - -
230 0 O 2 2 0 0 5
232 0 0 0 0 0 - -
233 l - - 1 l - -
222 l O 3 4 1 0 115
521 6 2 7 15 14 7 97
522 0 1 1 2 5 5 78
523 1 1 2 4 1 1 21
524 l O 6 7 2 0 52
543 1 1 8 10 1 1 124
544 ,1 0 5 6 2 O 51
545 1 1 1 3 l 1 114
273 0 2 3 5 0 7 5
274 1 0 5 6 1 O 39
201 .9 .5. .9 19 .9 .9 .99
Total 14 13 48 85 33 30 793

11“}!!! . —

63

Appendix 4. (continued) NUmber of Scotch pine trees attacked by the
European pine sawfly and the amount of attack per tree, June

 

 

 

 

 

1965.

MSFG No. Trees attacked Attack per tree_fl_
and Rose Rose
variety Kellogg Allegan Lake Sum Kellogg Allegan Lake
- - - - Number - - - - - - Colonies - - Per-
cent

rigensis
542 l 3 9 13 4 8 91
541 5 1 17 23 16 2 122
550 7 3 - 10 15 6 -
223 2 2 14 18 3 8 218
224 .1 .9 .9 19 _1 .9 .79
Total 16 9 49 74 39 24 507

armena

213 3 2 18 23 6 4 78
214 2 - - 2 6 - -
220 4 4 9 17 9 13 92
221 8 1 14 23 15 3 112
261 1 - 4 5 1 - 70
262 5 - - 5 7 - -
263 9 — - 9 18 - -
254 .9 ' .: _‘i .12 _:. _:.
Total 38 7 45 90 74 20 352

aquitana
212 5 - - 5 11 - -
238 5 7 ll 23 18 22 98
239 2 6 13 21 5 18 104
240 4 - - 4 2 - —
249 - - 7 7 - - 70
316 5 - - 5 12 - -
320 .9 - .1.1. 151. 11 _:. .99
Total 24 13 42 79 59 40 365

scotica
265 6 - - 6 20 - -
266 1 - - l 8 - -
267 7 - 10 17 10 - 159
268 .9 .: .: .fl. .9 .: ..:
Total 18 10 28 43 - 159

iberica
218 6 12 20 38 21 29 194
219 7 7 11 25 13 11 57
245 7 12 25 44 21 35 106
246 3 0 8 11 5 0 121
247 .9 19 .9. .99 .99 .11 .99
Total 32 43 73 148 80 92 546

64

Appendix 4. (continued) Number of Scotch pine trees attacked by the
European pine sawfly and the amount of attack per tree, June

 

 

 

 

1965.
MSFG No. Trees attacked Attack per tree
and Rose Rose
variety Kellogg, Allegan Lake Sum Kellogg Allegan Lake
- - - - Number - - - - - - Colonies - - Per-
cent
rhodopaea
243 2 8 9 19 4 24 44
244 7 8 l3 , 28 18 21 106
271 3 4 - 7 5 10 -
272 2 - 19 21 4 - 95
551 ..9. _‘ .: .2 .13 ' ..:
Total 23 20 41 84 43 33 245
illyrica
242 12 9 17 38 25 24 134
pannonica
552 7 - 20 27 40 - 140
553 .8. .z 99 91 99 19 199
Total 15 7 42 64 72 15 248
carpatica
314 3 - - 3 8 - -
borussica
202 5 4 18 27 20 12 89
219 19 1.1. 99 99 99 9.9 999
Total 24 15 43 82 73 36 368
hercynica
203 6 7 19 32 18 20 115
204 7 8 - 15 25 20 -
207 14 14 27 55 31 43 247
208 8 13 22 43 23 34 129
248 - - 18 18 - - 176
525 4 13 23 40 10 28 234
526 10 — - 10 24 - -
527 14 3 20 37 36 8 105
528 13 13 - 26 41 33 -
529 13 - - 13 39 - -
305 13 . 7 23 43 31 14 196
306 13 8 22 43 24 10 90
307 7 11 - 18 29 39 -
308 9 8 23 40 29 18 223
309 11 6 26 43 28 11 192
310 11 4 16 31 47 8 123
311 8 16 25 49 37 33 281

65

Appendix 4. (continued) Number of Scotch pine trees attacked by the
European pine sawfly and the amount of attack per tree, June

 

 

 

 

 

1965.
MSFG No. __~g Trees attacked Attack per tree
and Rose Rose
variety Kellogg Allegan Lake Sum Kellogg Allegan Lake
- - - - Number - - - - - - Colonies - - Per-
cent
312 12 8 22 42 27 21 227
313 7 - - 7 15 - -
315 15 - - 15 53 - -
319 ._-_ ..7. .99 .99 .: .19 15.4.
Total 195 146 309 650 567 350 2392
scotica X ?
269 12 - 29 41 62 - 260
270 .7. .: _:. .1 2.5. .: _:.
Total 19 29 48 87 - 260
polonica
211 10 14 25 49 31 32 246
317 .9 .8. 99 .99 99 99 12.9
Total 18 22 47 87 53 ‘54 369
haguenensis
206 ‘ 6 14 19 39 14 38 106
250 10 - 27 37 29 - 242
251 19 11 26 56 42 25 163
252 8 14 25 47 11 58 256
253 12 11 23 46 27 32 204
235 4 12 19 35 9 27 217
236 18 - - 18 88 - -
237 10 - - 10 31 - -
241 14 8 29 51 40 21 193
318 17 12 30 59 44 28 226
530 .19 19 .29 .2 .59 .99 -159
Total 135 94 227 456 385 259 1769
no described variety
554 4 - 9 13 9 - 69
555 4 - - 4 9 - -
556 6 - 17 23 23 - 62
557 6 - 12 18 17 - 38
225 4 7 25 36 26 16 200
205 4 - - 4 11 - -
209 .2 .: _:. ..9. .29 .; ..:
Total 37 7 63 107 123 16 189

66

 

 

 

 

 

 

Appendix 5. Trees with European pine sawfly damage.
Analysis of variance -- plantation 2—61

ecotype

source df SSQ MSQ F statistic
A variety

replicate

error

total
ecotype

source df SSQ MSQ F statistic
B variety

replicate

error

total
ecotype

source df SSQ MSQ F statistic
C variety 11 3.62 .329 1.87

replicate 7 1.82 .260 1.45

error 77 13.80 .179

total 95 19.24
ecotype

source df SSQ MSQ F statistic
D variety 3 1.34 .447 2.76

replicate 7 1.72 .246 1.52

error 21 3.41 .162

total 31 6.47
ecotype

source df SSQ MBQ F statistic
E variety 3 .12 .04 .13

replicate 7 .22 .03 .10

error 21 .63 .30

total 31 .97
ecotype

source df SSQ MSQ F statistic
F variety 1 .25 .25 .47

replicate 7 19.75 2.82 5.32*

error 7 3.75 .53

total

23.75

 

 

 

 

 

 

 

Appendix 5. (continued) Trees with European pine sawfly damage.
Analysis of variance -- plantation 2—61
ecotype
source df SSQ MSQ F statistic
G variety 13 27.44 2.11 1.51
replicate 7 15.56 2.22 1.58
error 91 127.06 1.40
total 111 170.06
ecotype
source df SSQ MSQ F statistic
H variety 8 29.26 3.66 2.00
replicate 7 7.21 1.03 .56
error 56 102.41 1.83
total 71 137.88
ecotype
source df SSQ MSQ F statistic
J variety 1 4.00 4.00 5.49
replicate 7 1.00 .14 .19
error 7 5.00 .71
total 15 10.00
ecotype
source df SSQ MSQ F statistic
K variety 4 3.35 .84 1.02
replicate 7 5.20 .74 .90
error 28 23.05 .82
total 39 31.60
ecotype
source df SSQ MSQ F statistic
M variety 1 .56 .56 .65
replicate 7 3.44 .49 .58
error 7 5.94 .85
total 15 9.94
ecotype
source df SSQ MSQ F statistic
N variety 4 3.40 .85 .95
replicate 7 8.00 1.14 1.28
error 28 25.00 .89

total

36.

4O

 

68

 

 

 

 

 

 

Appendix 5. (continued) Trees with European pine sawfly damage.
Analysis of variance -- plantation 11-61
ecotype
source df SSQ MBQ F statistic
A variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
B variety
replicate
error
total
ecotype
source df SSQ MSQ 1" statistic
C variety 11 2.29 .21 1.48
replicate 9 1.17 .13 .09
error 99 14.13 .14
total 119 17.59
ecotype
source df SSQ MSQ F statistic
D variety 3 .50 .17 1.00
replicate 9 2.10 .23 1.39
error 27 4.50 .17
total 39 7.10
ecotype
source df SSQ MSQ F statistic
E variety 3 .10 .03 1.00
replicate 9 .90 .10 3.03*
error 27 .90 .03
total 39 1.90
ecotype
source df SSQ MSQ F statistic
F variety 1 1.80 1.80 1.97
replicate 9 5.80 .64 .70
error 9 8.20 .91
total 19 15.80

 

 

 

 

 

 

 

Appendix 5. (continued) Trees with European pine sawfly damage.
Analysis of variance -- plantation 11-61
ecotype
source df SSQ MSQ F statistic
G variety 13 23.22 1.79 2.63**
replicate 9 9.99 1.11 1.63
error 117 80.21 .68
total 139 113.42
ecotype
source df SSQ MSQ F statistic
H variety 8 6.00 .75 .73
replicate 9 8.93 .99 .96
error 72 74.67 1.03
total 89 89.60
ecotype
source df SSQ ESQ F statistic
J variety 1 .45 .45 1.97
replicate 9 8.45 .94 4.12*
error 9 2.05 .23
total 19 10.95
ecotype
source df SSQ MSQ F statistic
K variety 4 2.92 .73 1.46
replicate 9 3.62 .40 .80
error 36 17.88 .50
total 49 24.42
ecotype
source df SSQ MSQ F statistic
M variety 1 .05 .05 .04
replicate 9 8.05 .89 .77
error 9 10.45 1.16
total 19 18.55
ecotype
source df SSQ MSQ F statistic
N variety 4 11.03 2.78 3.35*
replicate 9 10.82 1.20 1.44
error 36 30.17 .83
total 49 52.02

 

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Appendix 5. (continued) Trees with European pine sawfly damage.
Analysis of variance -- plantation 12-61
ecotype
source df SSQ MSQ F statistic
A variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
B variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
C variety 11 11.50 1.04 2.26*
replicate 7 9.00 1.28 2.78*
error 77 35.50 .46
total 95 56.00
ecotype
source df SSQ MSQ F statistic
D variety 3 5.84 1.95 1.19
replicate 7 7.72 1.10 .67
error 21 34.41 1.64
total 31 47.97
ecotype
source df SSQ MSQ F statistic
E variety 3 .68 .23 2.29
replicate 7 1.63 .23 2.35
error 21 2.07 .09
total 31 . 4.38
ecotype
source df SSQ MSQ F statistic
F variety 1 .56 .56 .30
replicate 7 5.44 .78 .42
error 7 12.94 1.85
total 15 18.94

 

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71

 

 

 

 

 

 

 

 

Appendix 5. (continued) Trees with European pine sawfly damage.
Analysis of variance -- plantation 12—61
ecotype
source df SSQ MSQ F statistic
G variety 13 15.79 1.21 1.34
replicate 7 47.49 6.78 7.53**
error 91 82.14 .90
total 111 145.42
ecotype
source df SSQ MSQ F statistic
H variety 8 13.52 1.69 2.58* g
replicate 7 31.78 4.54 6.93** 5
error 56 36.70 .65 E
total 71 82.00 ;
ecotype
source (if SSQ MSQ F statistic
J variety 1 .25 .25 .36 5
replicate 7 18.00 2.57 3.63 ;_
error 7 4.75 .69
total 15 23.00
ecotype
source (if SSQ MSQ F statistic
K variety 4 7.15 1.79 1.57
replicate 7 14.98 2.14 1.89
error 28 31.65 1.13
total 39 53.78
ecotype
source (if SSQ MSQ F statistic
M variety 1 .25 .25 .15
replicate 7 14.00 2.00 1.19
error 7 11.75 1.68
total 15 26.00
ecotype
source df SSQ MSQ F statistic
N variety 4 28.15 7.04 9.14**
replicate 7 20.18 2.88 3.74**
error 28 21.45 .77
total 39 69.78

 

 

 

 

 

 

 

Appendix 6. Branches defoliated by the European pine sawfly.
Analysis of variance -— plantation 2-61
ecotype
source df SSQ MSQ F statistic
A variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
B variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
C variety 11 21.08 1.92 1.38
replicate 7 26.50 3.78 2.72*
error 77 107.25 1.39
total 95 154.83
ecotype
source df SSQ MSQ F statistic
D variety 3 17.25 5.75 2.70
replicate 7 22.00 3.14 1.47
error 21 44.75 2.13
total 31 84.00
ecotype
source df SSQ MSQ F statistic
E variety 3 .38 .13 1.02
replicate 7 .88 .13 1.02
error 21 2.62 .12
total 31 3.88
ecotype
source df SSQ MSQ F statistic
F variety 1 5.60 5.60 .91
replicate 7 238.94 34.13 5.57**
error 7 42.90 6.13

total

287.44

 

 

I Illa—b..-

73

 

 

 

 

 

 

Appendix 6. (continued) Branches defoliated by the European pine
sawfly.
Analysis of variance -- plantation 2-61
ecotype
source df SSQ MSQ F statistic
G variety 13 205.68 15.82 .90
replicate 7 375.54 53.65 3.05**
error 91 1600.46 17.59
total 111 2181.68
ecotype
source df SSQ MSQ F statistic
H variety 8 182.76 22.84 1.22
replicate 7 259.27 37.04 1.99
error 56 1037.91 18.53
total 71 ”1479.94
ecotype
source df SSQ MSQ F statistic
J variety 1 16.00 16.00 5.59*
replicate 7 25.75 3.68 1.29
error 7 20.00 2.86
total 15 61.75
ecotype
source df SSQ MSQ F statistic
K variety 4 17.65 4.41 .96
replicate 7 18.80 2.69 .61
error 28 123.95 4.43
total 39 160.40
ecotype
source df SSQ MSQ F statistic
M variety 1 10.44 10.44 .50
replicate 7 124.44 17.78 .85
error 7 147.06 21.01
total 15 281.94
ecotype
- source df SSQ MSQ F statistic
N variety 4 24.50 6.15 .83
replicate 7 45.20 6.46 .87
error 28 206.30 7.37
total 39 276.00

 

 

 

 

 

 

 

Appendix 6. (continued) Branches defoliated by the European pine

sawfly.

Analysis of variance -- plantation 11-61

ecotype

source df SSQ MSQ F statistic
A variety

replicate

error

total
ecotype

source df SSQ MSQ F statistic
B variety

replicate

error

total
ecotype

source df SSQ MSQ F statistic
C variety 11 11.50 1.04 1.18

replicate 9 9.50 1.05 1.18

error 99 87.50 .88

total 119 108.50
ecotype

source df SSQ MSQ F statistic
D variety 3 5.10 1.70 .76

replicate 9 20.40 2.27 1.01

error 27 60.40 2.24

total 39 85.90
ecotype

source df SSQ MSQ F statistic
E variety 3 .10 .03 1.00

replicate 9 .90 .10 3.12*

error 27 .90 .03

total 39 1.90
ecotype

source df SSQ MSQ F statistic
F variety 1 5.00 5.00 .37

replicate 9 52.20 5.80 .43

error 9 121.00 13.44

total 19 178.20

 

 

 

 

 

 

 

Appendix 6. (continued) Branches defoliated by the European pine
sawfly.
Analysis of variance -- plantation 11-61
ecotype
source df SSQ MSQ F statistic
G variety 13 154.66 11.90 2.24*
replicate 9 62.57 6.95 1.31
error 117 621.63 5.31
total 139 838.86
ecotype
source df SSQ MSQ F statistic
H variety 8 137.76 17.22 1.85
replicate 9 84.46 9.38 1.01
error 72 670.24 9.31
total 89 892.46
ecotype
source df SSQ MSQ F statistic
J variety 1 .45 .45 .14
replicate 9 64.45 7.16 2.22
error 9 29.05 3.23
total 19 92.95
ecotype
source (if SSQ MSQ F statistic
K variety 4 36.60 9.15 1.82
replicate 9 30.90 3.43 .68
error 36 181.00 5.03
total 49 248.50
ecotype
source df SSQ MSQ F statistic
M variety 1 .80 .80 .00
replicate 9 87.00 9.67 .95
error 9 90.20 10.02
total 19 178.00
ecotype
source df SSQ MSQ F statistic
N variety 4 78.32 19.58 9.28**
replicate 9 60.32 6.70 3.17**
error 36 76.09 2.11
total 49 214.72

 

 

lliij

76

 

 

 

 

 

 

Appendix 6. (continued) Branches defoliated by the European pine
sawfly.
Analysis of variance -- plantation 12-61
ecotype
source df SSQ MSQ F statistic
A variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
B variety
replicate
error
total
ecotype
source df SSQ MSQ F statistic
C variety 11 8453.83 768.53 3.56**
replicate 7 8188.03 1169.72 5.14**
error 77 16638.10 216.08
total 95 33279.95
ecotype
source df SSQ MSQ F statistic
D variety 3 1775.34 5917.80 16.95**
replicate 7 2918.97 416.99 1.19
error 21 7329.91 349.04
total 31 12024.22
ecotype
source df SSQ MSQ F statistic
E variety 3 49.20 16.40 .43
replicate 7 73.10 10.44 .27
error 21 805.30 38.35
total 31 927.60
ecotype
source df SSQ MSQ F statistic
F variety 1 945.00 945.00 1.34
replicate 7 2346.88 335.27 .47
error 7 4940.50 705.78
total 15 8232.38

 

77

 

 

 

 

 

 

Appendix 6. (continued) Branches defoliated by the European pine
sawfly.
Analysis of variance -- plantation 12-61
ecotype
source df SSQ MSQ F statistic
G variety 13 7784.23 598.79 2.14*
replicate 7 13842.97 1977.57 7.08**
error 91 25427.91 279.42
total 111 47055.11
ecotype
source df SSQ MSQ F statistic
H variety 8 2538.69 317.34 1.02
replicate 7 10157.72 1451.10 4.69**
error 56 17337.53 309.60
total 71 30033.94
ecotype
source df SSQ MSQ F statistic
J variety 1 430.56 430.56 5.04
replicate 7 3455.90 493.70 5.77*
error 7 598.48 85.50
total 15 4484.94
ecotype
‘ source df SSQ MSQ F statistic
K variety 4 367.44 91.86 .48
replicate 7 2534.84 360.69 1.88
error 28 5366.16 191.65
total 39 8258.44
ecotype
source df SSQ MSQ F statistic
M variety 1 2.00 2.00 .00
replicate 7 1380.75 197.25 1.14
error 7 1215.00 173.57
total 15 2597.75
ecotype
source df SSQ MSQ F statistic
N variety 4 1470.35 367.59 1.18
replicate 7 1958.30 279.76 .90
error 28 8738.45 312.08
total 39 12167.10

 

 

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VITA

Name: William K. Randall
Place of Birth: West Branch, Michigan
Date of Birth: March 23, 1939

Education:
B. S., 1962
Michigan Technological University
Houghton, Michigan

M. S., 1966
Michigan State University
East Lansing, Michigan

Experience:
Firestone Plantations Company
Harbel, Liberia
West Africa
June 1962 - July 1964

 

U. S. Forest Service
Lake Wenatchee Ranger Station
Leavenworth, Washington
June 1961 - Sept. 1962

U. S. Forest Service
Mio, Michigan
Summers 1960, 1959, and 1958

Organizations:
Society of American Foresters
Xi Sigma Pi

MICHIGAN STATE UNIVERSITY LIBRARIES

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3 1193 03175 7796