V

the
By LARRY

WEBER

"W" magine yourself living in a globelike room with

greenish walls bulging outward and upward and
J L then arching in to meet above your head," the
naturalist Edwin Way Teale wrote. "Imagine such a room
constructed of succulent, edible material, forming a house
that at once provides food and shelter, plenty and
protection. That is what you would find if you
traded places with one of those gall
- - vw
insects that now

G O L D E N R O D G A L L S INDUCED BY THE G O L D E N R O O G A L L FLY

BY W A R R E N A B R A H A M S O N . B U C K N E L L UNIVERSITY

28

live in the
globular
swellings on the
stem of my hillside
goldenrods."
We have all seen these
curious swellings on goldenrod
stems, though usually we do not
imagine them as Teale did.
M

\INI II~DI

CI MIII R

2000

29

AN

INTRODUCTION

TO

GALLS

surprises inside

alls are
unique
growths on
plants, started in
reaction to the egg
laying or feeding of
gall-making insects
and mites. The
physical irritation
or chemical
A goldenrod gall fly larva
secretion of these
inside a goldenrod ball gall.
insect activities
results in a
distinctive enlargement or swelling of the plant,
ranging from simple to highly complex forms and
shapes. Each kind of gall maker can usually cause
only one kind of gall and prefers to use specific
plants.
Midges and other flies, gall wasps, and mites
make up the largest number of gall makers. Galls
are also caused by aphids, by sawflies, and less
often by a few kinds of moths and beetles,
nematodes, psyllids, and true bugs.
About 2,000 kinds of galls are known to exist in
North America on half the families of plants. They
Larry Weber is a teacher at Marshall School in Duluth. He
lives on an old farm in Carlton County, a great place to watch
critters.

30

Eastern s p r u c e
galls (right).
Hedgehog oak
gall (below).
Hackberry
nipple galls
(bottom).

and surprising outsides

may be on any part of a
plant, depending on the
gall maker's specialty:
| stems, twigs, leaves,
1 buds, flower heads,
fruits, and roots. They
5 resemble seeds, cones,
= burrs, flowers, balls,
| swellings, fur, and
| fingers.

Usually a particular
| insect forms a gall of a
distinctive shape on a
single species of plant,
such as the Canada goldenrod. Both woody and
nonwoody plants have
galls. Some are green,
others red or purple,
but most change color
with the seasons. Some
are seen only in
summer; some are
visible all year.

THE GOLDENROD

BALL

GALL

protected year-round

he goldenrod ball gall is a good choice to

study because scientists know about the life
cycle of the goldenrod gall fly, the insect that
causes it. In Minnesota this gall grows on two
species of goldenrodlate and Canada
goldenrodswhich are plentiful and easy to find in
almost any prairie, u n m o w e d field, or weedy place.
W A R R E N ABRAHAMSON. BUCKNELL UNIVERSITY

In late spring the female goldenrod

gall fly, only one-third of an inch
long, lays her eggs. She uses her
ovipositor, the pointed t u b e on the
end of her a b d o m e n , t o deposit eggs
on each tender goldenrod s h o o t .
Special cells on her feet help her
know this is the right kind of plant.
Only o n e gall usually a p p e a r s on
each stem.

WARREN ABRAHAMSON, BUCKNELL UNIVERSITY

Ten days later, the egg

hatches a n d the larva
burrows into the stem. As
the plant grows, the gall
begins t o thicken in
response t o the chewing
and secretions of the
larva. As it eats the nutritious gall tissue, it passes
through t w o larval stages.

32

W A R R E N ABRAHAMSON, BUCKNELL UNIVERSITY

T h e full-grown galls o f fall are

a b o u t 1 inch in d i a m e t e r .
Inside, t h e f a t larva is n o w fullsized t o o , a n d it is f o r m i n g a
tunnel t h a t s t o p s j u s t s h o r t o f
going all t h e way o u t s i d e . It
will use this tunnel in spring t o
escape. In t h e m e a n t i m e , it will
overwinter in t h e gall.

especially
from freezing
winter

Triggered by cold weather, the larva

makes a type of antifreeze called
glycerol. By keeping the liquid inside
the cells from freezing, the glycerol
protects the larva's delicate cell
m e m b r a n e s f r o m being destroyed
by ice crystals. T h e rest of the larva
can freeze solid. (See page 3 9 for a
winter project.)
After m a n y m o n t h s o f being
frozen a n d inactive, t h e larva
finally t u r n s into a p u p a a s t h e
w a r m spring w e a t h e r arrives. T w o

weeks later t h e a d u l t fly emerges,

s c o o t s t h r o u g h t h e tunnel, a n d
bursts t h r o u g h t h e thin gall wall.
T h e n t h e cycle begins again.

LARRY WEBER

33

THE GOLDENROD

BALL

GALL

Like other galls,

goldenrod galls
harbor many
different animals

GOLDENROD GALL FLY ADULT FEMALE

BY W A R R E N ABRAHAMSON

Plants are at the first level of

the center
of a busy
community

energy users. They collect energy

directly from sunlight and use it to
make sugars. For this reason they
are known as primary producers.
When a goldenrod produces a
gall, it provides specialized
nutrition with its inner gall tissue and shelter with its
tough outer layer. (In addition to supporting gall makers,
goldenrod feeds many other animals that eat everything
from the pollen to the roots.)

Gall makers are plant eaters, the second level of

energy users. Plant eaters are called the primary
consumers.The goldenrod gall fly larva feeds on the
goldenrod gall's plant tissue. The goldenrod gains
nothing from the larva inside it, while the larva
gains food and shelter. This one-sided relationship
is called parasitic.
Predators make up the third level of energy users in this
community. They prey on the gall-making larva.
One invader, the tumbling flower beetle, lays an egg in or near
the gall. Once hatched, the larva lives side by side with the fly
larva and will sometimes eat it. Animals that live in nests they do
not build are called inquilines.
Other parasitic invaders, called parasitoids, lay eggs right
inside the gall maker's eggs or larvae. One kind of
parasitoid wasp hatches inside the host larva, quickly eats
it, then feeds on the gall. Another kind of parasitoid wasp
hatches but stays inactive in the host larva through
summer. In fall, it forces the host larva to pupate early, then
takes over the gall to overwinter and finish growing.
Other predators include downy woodpeckers,
chickadees, and possibly gray squirrels. They peck and
chew their way into galls to devour the larva inside. Even ice
anglers sometimes open galls to use the larvae for bait.

TUMBLING FLOWER
BEETLE BY
PAI II UPLMRIRU

MORE

MINNESOTA

GALLS

large and
oak apple galls
O a k s a p p e a r t o be the m o s t p o p u l a r choice for t h e gall
makers of N o r t h America. M o s t o a k galls are c a u s e d by
small w a s p s called cynipids
o r gall w a s p s . A b o u t 8 0

p e r c e n t of the m o r e t h a n

7 0 0 different species of
gall w a s p s in the United
S t a t e s p r o d u c e galls
specifically on o a k trees.
M o s t n o t i c e a b l e of t h e
o a k galls p r o d u c e d by the
gall w a s p are t h e golfball-sized o a k a p p l e galls.
Oak apple galls form on
leaf stems or veins t h a t run
down the center of the leaf. Springtime oak apple galls look
like little green apples, thus accounting for their name. The
green galls blend with the leaves, but those that turn brown in
s u m m e r are easy to see.

blackberry knot galls

Perhaps the largest of Minnesota's galls are those f o u n d on
the blackberry plant. A m o n g the thorns on this brambly
bush is a solid, knotty, furrowed
swelling up to 6 inches long. Green

at first, the gall gets darker with age.

A g r o u p of larvae o f t h e tiny knot gall

w a s p causes these huge growths.

This w a s p is related t o those t h a t

form oak apple galls. On blackberries they live in a colony.

6
rrpfnpi

36

very small
jumping oak galls
W o u l d you be surprised if you s a w t h o u s a n d s o f tiny yellow
balls falling like rain f r o m o a k leaves, then j u m p i n g a r o u n d
on the ground?
T h e s e pinhead-sized galls a n d t h e j u m p i n g are c a u s e d by
j u m p i n g o a k gall w a s p larvae. T h e larvae's saliva c o n t a i n s
chemicals t h a t s t i m u l a t e t h e o a k leaf tissue t o f o r m yellow
blister-shaped galls.
As t h e galls m a t u r e , t h e y d e t a c h f r o m t h e leaf a n d rain
d o w n o n t o t h e g r o u n d . T h e galls j u m p a r o u n d until t h e y
b e c o m e l o d g e d in t h e soil, w h e r e t h e

WAYNE P ARMSTRONG. PALOMAR COLLEGE

w a s p s overwinter. At t i m e s t h e s e galls c a n
JEFFREY HAHN

b e c o m e s o n u m e r o u s t h a t they d i s c o l o r t h e o a k leaves
a n d even c a u s e t h e leaves t o fall early. Yet t h e trees stay
healthy.
J u m p i n g o a k galls are f o u n d in any o a k tree f o r only a year
o r t w o , usually d u r i n g times o f plentiful rain. So be sure t o
enjoy these marvels w h e n you find t h e m .

37

MORE

MINNESOTA

GALLS

cherry pouch galls

A mice forms cherry pouch galls, fingerlike projections on
t o p of cherry tree leaves. These numerous galls usually grow
in spring as the leaves grow. Their presence is a sign of mites
feeding and breeding. Anyone looking at cherry leaves during the s u m m e r might notice them. They d o n ' t occur on all
cherry trees, but they are a b u n d a n t when present. They
d r o p with the leaves, and the tree appears t o have never
been visited by these gall makers.

in all
shapes
and
textures

willow-cone galls

Even though these galls grow on willow shrubs, overlapping

scales make them look like pine cones. Sometimes a willow
has so many, it seems there's a gall at the end of every branch.
During the spring and early summer, a midge (a tiny gnatlike insect) lays eggs on new willow growth tips. T h e gall,
stimulated by the hatched larva, grows up to V/i inches long
and a b o u t 1 inch wide. Both the larva and the gall develop
during the warm m o n t h s . The larva b e c o m e s d o r m a n t in
winter and completes its growth
the following spring.
Many insects take advantage of
the n a p p i n g larva's large nest and
move in. S o m e of the uninvited
guests live as inquilines, others as
parasitoids. In one study,
researchers f o u n d 23 galls contained 5 6 4 insects, only 15 of
which were the original gall makers.
38

RONWINCH

GALL

PROJECTS

ow with the leaves off the trees, we can see

many of these galls and again get a sense of
the marvelous adaptability and diversity of
insects. Before the snow gets too deep, look for galls.
Cut open the different kinds of galls you collect and
try to find the gall makers. Certain kinds will contain
a larva inside; others will be empty.
Collect goldenrod galls in late fall or early winter
to observe the antifreeze effects of glycerol on
goldenrod gall fly larvae. Store harvested gall stems
outside, protected from rodents and mildew by a
cloth sack in a large, airy container.
When you are ready to begin, have a grown-up
help you crack open a frozen gall by twisting the tip
of a knife into it. You might
find a larva so hard it will
bounce. Put it into a dish
and observe it as it warms.
It will change from a frozen, |
white lump to a creamy, live |
wriggler. You can place the I
dish in a freezer for an hour |
or two to refreeze the larva.
Then watch it warm up all S
over again.

When you are finished

with this demonstration,

you might be able to get the larva to pupate. As a
larva, it must undergo obligate dormancy. So keep it in
the freezer until late spring. Then bring it to room
temperature. It should pupate within 24 hours, if
ready. Release the adult fly where you found the galls.
Here are a few gall web sices to explore:
http://www.naturenorth.com/winter/gallfly/Fgallfly.html
http://www.facstaff.bucknell.edu/abrahmsn/solidago/main.html
http://superior.carleton.ca/~kbstorey/insects.htm
http://daphne.palomar.edu/wayne/pldec97.htm
39