Impact of extrafloral nectar availability and plant
genotype on ant (Hymenoptera: Formicidae)
visitation to quaking aspen (Salicaceae)
Jonathon R. Newman, Diane Wagner,
1
Patricia Doak
Abstract—For quaking aspen (Populus tremuloides Michaux; Salicaceae) the rate of extrafloral (EF)
sugar secretion is increased by defoliation and decreased by drought. Although wholesale blocking of
EF nectar has been shown to reduce ant (Hymenoptera: Formicidae) visitation to aspen, the effect of
more subtle and realistic variations in nectar availability on ant recruitment is unknown. Working in
Alaskan boreal forest (United States of America), we reduced and supplemented EF nectar availability
on potted aspen ramets of three genotypes and surveyed visitation by free-living Formica fusca
(Linnaeus) (Hymenoptera: Formicidae). Ants were more responsive to a subtle increase in sugar
availability than to a decrease. While nectar reduction had no effect on ant visitation, nectar
supplementation increased ant visitation to one aspen genotype by 70% during an early summer trial.
Average ant visitation to different aspen genotypes varied during the late summer, indicating that aspen
genotype can influence attractiveness to ants. We conclude that natural induction of EF secretion in
response to herbivory may benefit aspen through improved ant recruitment, though the response is
dependent on aspen genotype and time of year. Differences among aspen genets in attractiveness to
ants could influence the relative success of genotypes, especially in settings in which aspen regenerates
from seed.
Introduction
Extrafloral (EF) nectar mediates defensive
mutualisms in many plant populations (Koptur
2005). Meta-analyses support that, in general, ant
visitation to plants expressing EF nectaries bene-
fits plants by reducing herbivory (Chamberlain
and Holland 2009; Rosumek et al. 2009: Trager
et al. 2010). The quantity and quality of food
rewards provided by the plant can affect the
strength of defensive mutualisms. For example,
ant visitation was positively associated with EF
nectar availability across several experimental
studies (e.g., Rudgers 2004; Rudgers and Strauss
2004; Kost and Heil 2005; Chamberlain and
Holland 2008), and an increase in EF nectar secre-
tion was associated with reduced leaf damage and
increased fruit set by lima bean (Phaseolus lunatus
Linnaeus; Fabaceae) (Heil 2004). Variation in the
rate of EF nectar secretion occurs in response to
environmental cues. Defoliation, application of
jasmonic acid, and exposure to volatile organic
compounds from damaged neighbours have been
shown to increase the rate of EF nectar secretion
across a broad range of EF nectary-bearing plant
species (Heil 2011), and drought stress has been
shown to reduce EF nectar secretion rate in a
smaller number of species (Yamawo et al. 2012;
Newman and Wagner 2013).
Quaking aspen (Populus tremuloides Michaux;
Salicaceae) expresses EF nectaries on a subset of its
leaves (Doak et al. 2007). Extrafloral nectaries
appear to function as an indirect defence against
herbivory for boreal aspen, as EF nectary expres-
sion by ramets is negatively related to damage by
the leaf miner Phyllocnistis populiella Chambers
(Lepidoptera: Gracillariidae) (Mortensen et al.
2011). EF nectar secretion by aspen plants varies in
response to environmental stimuli. In a greenhouse
experiment, 40% defoliation increased the EF sugar
secretion rate by about 25%, whereas drought
decreased the secretion rate of some aspen geno-
types by a similar magnitude (Newman and
Wagner 2013).
J.R. Newman, D. Wagner,
1
P. Doak, Institute of Arctic Biology and Department of Biology and Wildlife,
University of Alaska Fairbanks, Fairbanks, Alaska 99775, United States of America
1
Corresponding author (e-mail: diane.wagner@alaska.edu).
Subject editor: Justin Schmidt
doi:10.4039/tce.2015.38
Received 19 August 2014. Accepted 26 February 2015.
Can. Entomol. 00:1–7 (2015) © 2015 Entomological Society of Canada
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