Indirect Benefits of Scale Insects

for Ant-Defended Trees

Elizabeth Pringle et al.

Photo 1. A Cordia alliodora tree in flower on a hill overlooking the forest in the Area de
Conservación Guanacaste, Sector Santa Rosa, Costa Rica, in the dry season.Volcanoes are visible in
the distance. Photo by Elizabeth G. Pringle.

The Neotropical tree Cordia alliodora produces hollow stem nodes that Azteca ant colonies use as
housing. These ants patrol and defend the tree from leaf-eating herbivores. Ants also tend tree-sap-
feeding scale insects inside stem nodes, and these scale insects produce sugar-rich excretions that ants
consume. Experiments and observations showed that: larger ant colonies were associated with more
scale insects and were better leaf defenders than smaller colonies; individual ants that ate more sugar
were more aggressive defenders; and scale insects were more numerous in the trees’ young tissues,
which appeared to stimulate better ant defense of young leaves. These findings suggest that although
scale insects are herbivores and therefore potentially costly to trees, they also indirectly benefit trees
by increasing the effectiveness of ant leaf defense.These indirect benefits point to positive feedback be-
tween tree investments and ant services, which may stabilize ant–tree mutualisms in evolutionary time.

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Photo 2. Hollow swellings at branch

nodes of the tree C. alliodora, known as
domatia, house ant colonies and scale
insects. Visible here are scale insects of
the family Coccidae, feeding on tree sap
through the domatium walls, as well as
Azteca pittieri ant larvae. Photo by Jeffrey C.
Miller.

Photo 3. (Left) Leaves of C. alliodora that have experienced low herbivory in the Chamela-
Cuixmala Biosphere Reserve, Jalisco, Mexico. (Right) Leaves of C. alliodora that have been highly
herbivorized by beetle larvae, leaf-mining Lepidoptera, and caterpillars in the Area de Conservación
Guanacaste, Costa Rica. Photos by Elizabeth G. Pringle.

102 Bulletin of the Ecological Society of America

 Photo 4. A noctuid moth caterpillar (Cropia templada) chows down on a C. alliodora leaf,
eating from the tip of the leaf towards the base. Photo by Elizabeth G. Pringle.

Photo 5. A C. templada caterpillar drops from a leaf by a silk thread after being bitten
repeatedly by Azteca ants. Photo by Arpita Sinha.

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Photo 6. Azteca pittieri ants drinking the 70% sugar solution in the lab experiment.
Ants were fed with either 70% or 2% sugar diets for three weeks, and were provided with
unlimited protein (tuna) and water. Ants fed the 70% sugar diet were much more likely than
those fed the 2% sugar diet to attack a caterpillar herbivore placed in the container. Photo
by Elizabeth G. Pringle.

104 Bulletin of the Ecological Society of America

 Photo 7. Cordia alliodora domatia arranged from oldest (left) to youngest (right). Ants
enter domatia when they are young, soft, and green, and domatia become woodier with age.
Young leaves are associated with young domatia at the end of growing shoots. Scale insects
reached higher densities in younger domatia, which may encourage better ant defense of
young leaves. Photo by Elizabeth G. Pringle.

These photographs illustrate the article “Indirect benefits of symbiotic coccoids for an
ant-defended myrmecophytic tree,” by Elizabeth G. Pringle, Rodolfo Dirzo, and Deborah M.
Gordon, which will appear in Ecology 91(1):37–46, January 2011.

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