The Red pine scale in North America : a report to the 1975 Eastern Plant Board

John F. Anderson
The Red Pine Scale in North America

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ronnecticut Libraries

GAVLORD RG

BULLETIN 765

JULY 1976

A Report to the 1975 Eastern Plant Board

THE RED PINE SCALE
IN NORTH AMERICA

John F. Anderson

Chairman

Robert P. Ford

John D. Kegg

John H. Risley

siveo

UNIVERSITY LIBRARY

STAiJ

THE CONNECTICUT AGRICULTURAL EXPERIMENT STATION. NEW HAVEN

\

Members of the 1974 Eastern Plant Board appointed a four-man committee to make an in-
depth evaluation of the present and future status of the red pine scale, Matsiicoccus resinosae
B. & G., in North America. Herewith is our report.

John F. Anderson, Chalrmarv

Department of Entomology

The Connecticut Agricultural Experiment Station

New Haven 06504

Robert P. Ford

USDA Forest Service, Northeastern Area
State and Private Forestry
Portsmouth, N.H. 03801

John D. Kegg

Division of Plant Industry

New Jersey Department of Agriculture

Trenton 08625

John H. Risley

Division of Lands and Forest

New York State Department of Environmental Conservation

Sherburne 13460

THE RED PINE SCALE IN NORTH AMERICA

John F. Anderson, Robert P. Ford, John D. Kegg, and John H. Risley

The red pine scale was initially found in 1946 in
red pine plantations in western Connecticut (Plumb
1950). The infestation was extensive, indicating that
the insect had b^en present for several years prior to
its recognition. It was recognized as an undescribed
species of Matsucoccus and subsequently described
and named Matsucoccus resinosae by Bean and God-
wan ( 1955 ) . It now occurs in Connecticut, New York,
and New Jersey. This species is presently estabhshed
south of the natural range of red pine, but it is mov-
ing north and westward and may, in time, infest nat-
ural stands. Inasmuch as there are no suitable pro-
cedures for coping with this pest, which kills its host,
this insect must be considered the most serious pest
of red pine.

TAXONOMY

Red Pine

( Harlow and Harrar 1958,
Critchfield and Little 1966 ) .

Order • Coniferales

Family • Pinaceae

Genus • Pinus

Subgenus * Pinus

Section • Pinus

Subsection • Sylvestres

Species • resinosa Ait.

Red Pine Scale

( Bean and Godwin 1955 )

Order

•

Homoptera

Superfamily

«

Coccoidea

Family

•

Margarodidae

Tribe

•

Matsucoccini

Genus

•

Matsucoccus

Species

•

resinosae B. & G.

BIONOMICS

Red Pine

The silvics of red pine have been discussed by
Powells (1965). It ranges through southern Canada
from Newfoundland to southeastern Manitoba and
south through the Lake states and northeastern U.S.,
with outliers in Illinois and West Virginia. The na-
tural stands of red pine are generally confined to
sandy soils of low fertility, although it is often found
in small groups near swamps and on rocky cliffs.
Heavy cutting in the early 1800s depleted many nat-
ural stands, which did not reproduce because of
competition from more aggressive species such as
hemlock, white pine, and hardwoods. The largest
stands now occur in Quebec, Ontario, New York,
Minnesota, Michigan, and Wisconsin (Table 1).

Red Pine Scale

Life Cycle. The life cycle has been discussed in de-
tail by Bean and Godwin (1955) and Duda (1961).
The stages include the egg, first stage larva, inter-
mediate stage, and adult.

(a) Egg. These hatch in 15-20 days in the labora-
tory. Adults of the overwintering generation lay eggs
from May until the first week of July. Adults of the
summer generation lay eggs from about the middle
of August until the last of October. Eggs are laid in
ovisacs in cracks or crevices in the bark and axils of
branches, under loose bark scales, and on needles.

(b) First stage larva. This is the overwintering
stage. Duration varies from 45-50 days for the sum-
mer generation and from 235-260 days for the over-
wintering generation. The newly hatched larvae are
active and may crawl considerable distances before
settling down to feed. Suitable feeding sites are un-
der loose bark scales or in cracks and crevices where

Connecticut Agricultural Experiment Station

Bulletin 765

the crawlers have access to the succulent inner bark.
SessCe or motile forms of this stage may be found
throughout the year.

(c) Intermediate stage. This is a sessile feeding
stage. It remains in the same location selected by the
first stage larva. Intermediate stages of the overwin-
tering generation occur in the latter part of April into
early July. Those of the summer generation occur
from mid July into October.

(d) Adult. The male forms that emerge from the
cast skins of the intermediate stages are not true
adults but pre-adults. Almost immediately after emer-
gence, these pre-adult males spin a loose, fluffy,
spindleshaped cocoon of wa.xy threads whereupon
transformation to the true-adult stage occurs. Females,
which do not pupate, are wingless and resemble the
pre-adult males but are much larger. They crawl over
the bark in search of suitable oviposition sites. Males
are winged and locate females via a pheromone
(Doane 1966). Females lay an average of 262 eggs.

(e) Generations. There are two generations per
year and sometimes there is a partial third generation
(Bean, personal communication).

Dispersal. Dispersal mechanisms are not thoroughly
understood. Females are wingless, accounting in part

Table 1. Red Pine Round wood Resource in North America

million cubic

foot volume

Location

Acres of Plantations'

(all stands)'

USA

Connecticut

9,900

58.7

Maine

15,700

42.5

Massachusetts

7,400

23.3

Minnesota

199,000

199.5

Michigan

723,500

289.6

Ohio

33,900

16.3

Pennsylvania

32,200

NA*

Rhode Island

600

11.8

New Hampshire

16,000

49.1

New Jersey

9,500"

NSV=

New York

95,900

327.3

Vermont

15,200

32.9

Wisconsin

239,400

241.1

West Virginia

200

NSV

Indiana

1,300

NSV

Iowa

1,800

NSV

Canada'

New Brunswick

45.0

Nova Scotia

26.0

Ontario

750.0

Quebec

222.0

TOTAL

2335.1

'Data obtained from Kingsley and Mayer ( 1972).

'Data obtained from the most recent forest survey by the U.S.

Forest Service.

'Data obtained from Horton and Bedell ( 1960).

'NA — Not available.

''NSV — No significant volume.

"Data from John D. Kegg.

for the slow spread of this insect (Doane 1959a).
Also, red pine is sparsely scattered throughout much
of the infested area (Mclntyre 1956). Presumably,
natural dispersal occurs only during the motile stages;
i.e. first stage larva and female. First stage larvae may
become air-borne and have been recorded to travel a
c^uarter of a mile (Bean and Godwin 1955). Birds
and mammals may also transport the motile stages
(Duda 1961). Motor vehicles are also known to trans-
port these insects, and this means of transport may
account, in part, for infestations occurring initially
along roadsides (Duda 1961). All stages may be
carried on nursery stock and timber and other red
pine products that are transported out of infested
areas. Wind, however, is probably the natural and
most important means of dispersal.

Natural mortality factors. Life table studies suggest
that the primary mortality factor is the degeneration
of phloem cells following repeated attacks on trees
(Duda 1961). This effect on the host prevents the
establishment of first stage larvae. Predators are also
considered important, but obviously, these biological
agents are not sufficient to protect plantations from
being heavily infested. Predators include Xenotra-
cheliella inimica Drake and Harris (Hemiptera: An-
thocoridae), a cecidomyid that preyed upon eggs,
Chrysopa sp. nr. cornea, Hemerobius stigmatus Fitch,
Mulsantina picta (Rand), Crematogaster cerasi Fitch,
and mites belonging to die genus Amjstis, along with
several species of spiders and nuthatches (Duda 1961,
Bean and Godwin 1955, Plumb 1950). No insect
pathogens or parasitoids have been reported.

Hartzell ( 1957 ) reported that first stage larvae were
highly susceptible to temperatures of — 10°F. Few
survived two exposures of —10° for four hours. This
susceptibility may, in part, explain the slow north-
ward spread of this insect.

Hosts. Aside from red pine, M. resinosae has been
reported infesting three species of exotic pines in
North America. Infestations have been reported on
Chinese pine (P. tabulaeformis Carr. ), Japanese red
pine (P. densiflora Sieb. and Zucc. ) and Japanese
black pine (P. thunbergiana Franco) (Hartzell 1957,
Duda 1962, 1964b).

Red pine scale was reported not to attack any of
the following eighteen species of pine (Hartzell
1957):

Pinus ari.'itata Engelm.

P. armandi Franch.

P. ayacahuite Ehrenb. (P. hamata Roezl.)

P. hungeana Zucc.

P. cembra L. var. sibirica Loud.

P. flexilis James

P. griff ithii McClelland (P. exceha Wall.)

The Red Pine Scale in North America

P. heldreichii Christ, var. leucodermis (Ant.) Markgraf ex

Fitschen.
P. koraiensis Sieb. & Zucc.
P. lamhertiana Dougl.
P. monticola Dougl.
P. mugo Turra var. rostrata Hoopes (P. montana Mill., P.

uncinata Ramond.)
P. nigra poiretiana (Ant.) Aschers. & Graebn. (P. laricio

Poir. var. calahrica Loud. )
P. parviflora Sieb. & Zucc. (P. pentaphylla Mayr.)
P. pence Griseb.
P. ponderosa Dougl.
P. rigida MUl.
P. sylvestris L. var. rigensis Loud.

Survival in cut logs. Infested trees have been and
are continuing to be salvaged and are often shipped
outside of infested areas. These salvage operations
are often carried out during the wanter, and the ques-
tion arises as to whether the first stage larvae can
survive to adulthood the following spring on cut logs.
Data by Duda (1964a) suggest that survival does oc-
cur and that logs could carry this species unless they
are debarked or otherwise utilized in such a fashion
that would destroy the overwintering larvae.

Geographical distribution. Red pine scale occurs in
Connecticut, New York, and New Jersey. The in-
fested areas within these states are shown in Figure 1.
In Connecticut, infestations have been moving north
and eastward at a rate of about 1 or 2 miles per year,
though in recent years the spread may be more rapid.
This scale presently occurs throughout much of Fair-
field and New Haven Counties, southern Litchfield
County, southern portions of Hartford County, a
western portion of Middlesex County, and one loca-
tion in New London County.

The initial infestation in New Jersey was found
near the Wanaque Reservoir in Passaic County. All
infestations to date have occurred within 10 miles of
the initial infestation. Infestations also occur in Ber-
gen County.

In New York, red pine scale was initially found in
1950 in East Norwich, Long Island, and in lower
Westchester County. Spread has been at 1 to 5 miles
per year. The scale now occurs in all but the north-
western part of Westchester County, on most of Long
Island, in southeastern Putnam County, the extreme
southeastern tip of Orange County and the extreme
southwestern tip of Rockland County.

RELATED SPECIES AND THEIR
GEOGRAPHICAL DISTRIBUTION

Red Pine

The following species all belong to the subsection
Sylvestres and are closely related to P. resinosae
(Critchfield and Little 1966) :

P. tropicalis Morelet, Tropical pine. Isle of Pines and western
Cuba.

P. nigra Arnold, Austrian pine. Ranges widely through south-
em Europe from Spain to Austria and throughout much of the
Balkan Peninsula east to southern Russia and south to Turkey,
on the islands of Cyprus, Sicily, and Corsica, with outliers in
Algeria and Morocco.

P. heldreichii Krist, Heldreich pine. Balkan Peninsula and
southern Italy.

P. mugo Turra, Swiss mountain pine. Central and southern
Europe, ranging east to the USSR, south to Rumania, Bulgaria
and central Italy, north to Germany and Poland, and west to
eastern Spain.

P. pinaster Ait., Maritime pine. Ranges from the Iberian
Peninsula to southern France, to the west coast of Italy, in
northern Africa from Morocco to Tunisia, and to the islands of
Corsica and Sardinia.

P. halepensis Mill., Aleppo pine. Widely distributed in the
Mediterranean region.

P. hrutia Ten. The eastern Aegean region, Turkey, the
Levant, on the Mediterranean islands of Crete and Cyprus, and
in the Black Sea region.

P. sijlvestiis L., Scotch pine. This is the most widely dis-
tributed species of pine and grows throughout northern Eurasia,
from Scotland, southern Spain, northern Greece and northern
Turkey to northern Manchuria, and the Sea of Okhotsk.

P. densiflora Sieb. & Zucc, Japanese red pine. Japan, Korea,
to eastern Manchuria and adjacent USSR, and the Shantung
Peninsula of China. Its distribution on the mainland of Asia is
uncertain.

P. thunbergiana Franco, Japanese black pine. Japan and
southern Korea.

P. massoniana Lamb., Masson pine. Widely distributed in
China.

P. taiwanensis Hayata, Taiwan red pine. Taiwan.

P. hichuensis Mayr, Luchu pine. Ryukyu Islands between
Japan and Taiwan.

P. htvangshanensis Hsia, Hwangshan pine. Central China.

P. tahulaeformis Carr., Chinese pine. Occupies a broad dis-
continuous belt in China.

P. tjunnanensis Franch., Yunnan pine. Confined to Yunnan,
southern Szechuan, and the western parts of Kwangsi and
Kweichow.

P. insidaris Endl., Khasi pine. Southeast Asia.

P. merkusii Jungh and deVriese, Merkus pine. Ranges from
eastern Burma, northern Laos, northern Viet Nam and the
island of Hainan, south to Cambodia, southern Viet Nam and
the island of Sumatra, and east to the Philippine Islands.

Red Pine Scale

The following species also belong to the genus
Matsucoccus (Bean and Godwin 1955, Duda 1961):

Species of

Matsucoccus

Host

Nearctic

Distribution

acalyptus Herbert

Pinus edulis

California, Arizona,

Engelm.

New Mexico, Utah,

P. monophylla

Colorado, and Idaho

T. & F.

P. balfouriana

G. & B.

P. aristata

Engelm.

alahamae Morr.

Pinus sp.

Alabama

bisetosus Morr.

P. ponderosa

California, Oregon,

Laws.

and Arizona

P. jeffreyi

G. &B.

Connecticut Agricultural Experiment Station

Bulletin 765

Species of

Matsucoccus

Host

Nearctic (cont.)

P. sabiniana

Dougl.
P. radiata

D. Don.
P. contorta

Dougl,

Distribution

californicus Morr.

P. ponderosa
P. jeffreyi

California and
Arizona

degeneratus Morr.

P. ponderosa

Arizona

eduli Morr.

P. edulis

Arizona

fasciulensis Herbert

P. ponderosa

California and

P. jeffreyi

Oregon

P. sabiniana

gallicolus Morr.

P. rigida

New England west

Mill.

to Ohio and

P. echinata

Mis.souri and south

Mill.

to Florida and

P. ponderosa

Georgia

P. taeda L.

P. virginiana

Mill.

P. glabra Walt.

P. serotina

Michx.

monophyllae McK.

P. monophylla
P. edulis

California

paucicicatrices

P. lamhertiana

California, Oregon,

Morr.

Doug.

Montana, and

P. monticola

Wyoming

Doug.

P. flexilis

James

secretus Morr.

P. ponderosa

California, Nevada,
Arizona. New
Mexico, and
Colorado

subdegeneratus

P. occidentalis

Dominican Republic

Morr.

(cubensis) Griseb.

vexillorum Morr.

P. ponderosa

California, Nevada,
Arizona, New
Mexico, and
Colorado

macrocicatnces

P. strobus L.

Ontario, Quebec,

Richards

New Brunswick,
Nova Scotia, and
New Hampshire

Palearctic & Oriental

M. boratynskii

Pinus sp.

Russia

Borkhsenius

M. feytaudi

P. maritima

France

Ducasse

Mill.

M. josephi

P. halepensis

Israel

Bodenheimer &

Mill.

Harpaz

M. pini (Green)

P. sylvestris L.

England

M. matsumurae

P. thunbergiana

Japan

( Kuwana )

Franco

M. sinensis Chen

P. yunnanensis
Franch.

China

M. yunnanensis
Ferris

P. yunnanensis
Franch.

China

ORIGIN OF RED PINE SCALE

The origin is unknown. The suggestion was made
that it was brought into this country on pines planted
on the grounds of the World Fair in 1937 and then
carried to Connecticut (Doane 1959b).

Although its origin is unknown, the evidence sug-
gests tliat it is indeed an imported pest from Eurasia.
Red Pine is the only species of the typically Eurasian
group of pines in the subsection Sylvestres that is na-
tive to North America (Critchfield and Little 1966).
The host range of M. resinosae is unknown, but aside
from red pine, it does attack three species of pine, P.
densiflora, thunbergiana, and tahulaeformis that are
native to Asia. It is not known to infest species of pine
that occur naturally in Europe. These data suggest
that red pine scale may have originated on a species
of pine native to Asia.

EFFECT ON HOST

Infested red pine, including mature trees, eventual-
ly die from the feeding of the scale (Bean and God-
win 1955, Duda 1961, Doane 1959b). Trees often die
quickly with little stunting or change in form (Duda
1961). The progressive morphological and histological
effects on the tree have been described (Bean and
Godwin 1955, Duda 1961). Plumb (1950) suggested
that death may, in part, be caused by a toxin pro-
duced by the scale.

Japanese red pine has succumbed to scale attack
in New York.

CHEMICAL CONTROL

No methods are available to protect plantations
from succumbing to scale attack. Ornamental pines
may be protected by spraying repeatedly with a 2%
oil emulsion (Bean and Godwin 1955). Weekly ap-
plications of Cygon and malathion against motile first
stage larvae are also effective, though neither of these
two insecticides is registered for this use (Duda, per-
sonal communication).

POTENTIAL IMPACT OF RED PINE SCALE

M. resinosae is attacking trees outside the natural
range of red pine. Should the scale extend into the
natural range of its host, red pine would become a
rare, rather than an uncommon species. It is suspected
that the red pine in stands would be killed, but some,
if not many, isolated trees might survive. Reproduc-
tion of natural stands would be unlikely because of
unsatisfactory seedbed and sunlight.

Red pine is a relatively unimportant species in
terms of the total forest of North America. It supplies
a very small fraction of the timber harvested when
compared to white pine, southern pines, Douglas fir,
and Ponderosa pine. The chief value of red pine Hes

The Red Pine Scale in North America

in its adaptability to reforest on dry sites, its aesthetic
aspects, and its use as pulp, posts and poles. The
popularity of red pine for reforestation is evident by
the many plantations ( Table 1 ) ( Kingsley and Mayer
1972).

Potential losses of red pine are in the millions of
cubic feet of both natural and planted stock. The
total red pine round wood that is threatened is shown
in Table 1.

Damage to red pine cannot be measured in saw
timber or pulpwood values alone. The impact on
aesthetic values is illustrated in New Jersey where in
a single year 156 ornamental red pines were cut and
burned because they were infested (Kegg 1974). If
these trees were replaced, the cost would average
$150 per tree.

PRESENT STATE POLICIES

Connecticut guidelines are as follows:

(1) Shippers of red pine timber are encouraged to
move their products during the cold months.

(2) Movement of red pine at any time within
Connecticut from one uninfested location to another
is permissible.

(3) Movement of red pine at any time during the
year from one location to another within the infested
area is permissible.

(4) Nurserymen within infested areas are en-
couraged to destroy their plantings.

The New Jersey State Board of Agriculture declared
the red pine scale a "dangerously injurious insect" in
1961, and efforts were initiated to remove and destroy
all known infestations. This policy continues to the
present. The purpose of destroying infested trees is to
reduce spreading the insect to pines in other areas of
the state. To date, 331.5 acres of plantation and over
1,000 ornamental red pines have been cut and des-
troyed.

New York has no quarantine on the red pine scale.
Some eradication was attempted to 1969, but this
pohcy has been discontinued.

CONCLUSIONS

(1) Biological control. Scale insects have been the
targets of numerous successful biological control at-
tempts (DeBach et al. 1971). In fact, the proportion
of success has been higher in the coccoideae than in
any other group of pests.

M. resinosae may be found in Eurasia. Biologi-
cal control agents could then be shipped to the
United States for colonization. Efforts could also
be expended to colonize on M. resinosae natural
enemies isolated from other Matsucoccus species
in North America and Eurasia. Several successful
biological controls, using enemies of allied species,
have been reported (Pimentel 1963). This latter ap-

proach would of necessity require a thorough study
of the bionomics of the genus Matsucoccus.

(2) Regulatory measures. Regulations that could
be used to limit the spread of this insect need ex-
amination. A quarantine was rejected by the Eastern
Plant Board in 1965 because the scale had already
spread considerably. Nevertheless, the State of New
Jersey does quarantine and it would seem that some
regulatory measures might impede the progress of this
insect at least until satisfactory methods of coping
with it have been developed. Two suggestions follow:

a. Determine the feasibility of removing major
scale infestations around parking lots, picnic and
recreational areas and along major highways.

b. Determine the feasibility of establishing a
barrier zone with the assistance of federal funds.

(3) Dispersal. Intensive experiments on disper-
sal mechanisms of the red pine scale seem worth-
while.

(4) Weather. The role of low winter tempera-
tures in limiting the spread of this species into the
natural red pine growing areas of North America
needs re-examination.

(5) Host. The mechanisms by which the scale
ultimately kills the tree need investigation.

(6) Chemical control. Methods of controlling
scales on ornamental trees with chemicals need to
be re-examined and thoroughly studied.

(7) Early detection. Methods of early detection
of infestations need to be developed.

ACKNOWLEDGMENTS

We express our appreciation to the following individuals who
contributed to the report:

Charles C. Doane, George R. Stephens, and George W.
Schuessler, The Connecticut Agricultural Experiment Station.

Jim Bean and Neal Kingsley, U.S. Forest Service

Edward J. Duda, University of Connecticut

E. G. TerriU, New York State Department of Environmental
Conservation

Douglas R. Miller, U.S. Department of Agriculture

Connecticut Agricultural Experiment Station

Bulletin 765

REFERENCES CITED

Bean, J. L. and P. A. Godwin. 1955. Description and bionomics
of a new red pine scale, Matsvcoccus resinosae. For. Sci.
1: 164-176.

Critchfield, W. B. and E. L. Little, Jr. 1966. Geographic dis-
tribution of the pines of the world. USDA For. Ser. Misc.
Publ. 991: 97 p.

DeBach, P., D. Bosen and C. E. Kennett. 1971. Biological con-
trol of coccids by introduced natural enemies. In Biological
Control, C. B. Huffaker (ed.). pp. 165-194.

Doane, C. C. 1959a. The red pine scale, Matsucoccus resinosae.
Conn. Woodlands 24( 1 ) : 8-9.

. 1959b. The red pine scale. Conn. Agric. Expt. Sta.

Circ. 207: 1-7.

. 1966. Evidence for a se.\ attractant in females of the

red pine scale. J. Econ. Entomol. 59: 1539-40.

Duda, E. J. 1961. Some aspects of the biology and ecology of
the red pine scale, Matsucoccus resinosae B. & G.
(Homoptera: Margarodidae). Univ. Mass. Ph.D. Thesis.
168 p.

. 1962. The genus Matsucoccus with special reference

to M. resinosae B. & G. Scientific Tree Topics 2: 1-10.

. 1964a. Survival of the red pine scale on cut logs.

Ibid. 2(10): 3,4.

.1964b. An infestation of Matsucoccus resinosae B. & G.

on Pinus thunbergii Pari, Ibid. 2( 10) : 9.

Fowells, H, A. (ed.). 1965. Silvics of forest trees of the United
States. USDA-FS Agricultural Handbook 271 : 762 p.

Harlow, W. M. and E. S. Harrar, 1958. Te.\tbook of Den-
drology. McGraw-Hill, New York. 561 p.

Hartzell, A. 1957. Bed pine scale with special reference to its
host plants and coldhardiness. Contrib. Boyce Thompson
Inst. 18: 421-28.

Horton, K. W. and G.H.D. Bedell. 1960. White and red pine.
Can. Dept. North. Affairs & Nat. Res. Forestry Branch
Bull. 124.

Kegg, J. D. 1974. Red pine scale status. Unpubhshed report.
N.J. Dept. Agr., Trenton, N.J. 1 p.

King.sley, N. P. and C. E. Mayer. 1972. The e.xtent, com-
position, and general condition of Northern conifer plan-
tations in the Northeast region. Unpublished paper
presented at the Conference on the Utilization of Planta-
tion-grown Northern Softwoods, Syracuse, New York.

Mclntyre, T. 1956. Matsucoccus resinosae B. & G. on red pine
in Connecticut. lOth Intern. Congr. Entomol. Proc.
(Montreal) 4:. 369-371.

Pimentel, D. 1963. Introducing parasites and predators to con-
trol native pests. Canadian Entomol. 95: 785-92.

Plumb, G. H. 1950. A new and serious insect pest of red pine.
Conn. Agric. E.xpt. Sta. Special Circ. 4 p.

Figure I. Shaded areas show infested areas of red pine scale In Connecticut, New York, New Jersey and Long Island.

University of
Connecticut

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