678 Invasive Stink Bugs and Related Species (Pentatomoidea)
15.1 Introduction
15.1.1 Scope: Families in Hemiptera: Heteroptera
We review here semiochemicals identified from species in the true bug superfamily Pentatomoidea,
which includes among others the families Pentatomidae, Acanthosomatidae, Plataspidae, Scutelleridae,
Cydnidae, and Tessaratomidae. This does not include some well-studied groups in the infraorder
Pentatomomorpha, such as Coreoidea (Alydidae, Coreidae, and Rhopalidae), Lygaeoidea (Berytidae,
Blissidae, Cymidae, Geocoridae, and Lygaeidae), Aradidae, and Pyrrhocoridae. Subfamilies and tribes
of Pentatomidae are classified according to Rider (2015).
15.1.2 Scope: Definitions of Semiochemicals
Semiochemicals identified from the Pentatomoidea, as with other organisms, fulfill a wide range of com-
munication roles for senders and receivers (terminology from Nordlund and Lewis 1976). Pheromones
are used for communication between members of the same species and may be classified as sex phero-
mones, aggregation pheromones, alarm pheromones, and other categories, according to their functional
roles. We cover these in detail in Section 15.2. Allomones (Section 15.3), which the common name “stink
bug” connotes, are repellent or repugnant chemicals used against members of other species such as
predators and parasitoids; as such, they benefit the sender at the expense of the receiver. Conversely,
kairomones benefit the receiver at the expense of the sender, and most examples discussed below involve
the exploitation of pheromonal or allomonal signals by natural enemies whose hosts or prey are pentato-
moid bugs, as summarized in Section 15.4. Plant volatiles that are released in response to pentatomoid
herbivory are considered synomones when they attract natural enemies of phytophagous bugs, thus con-
ferring a benefit to the plant which produces them, in helping to protect it from herbivory (Section 15.4.5).
In addition, several stink bug aggregation pheromones attract other species of stink bugs, which could
be considered a kairomonal or synomonal interaction, depending on whether the sender experiences fit-
ness losses or gains, respectively (Section 15.5.1.1). From the above statements it is apparent that a single
semiochemical, or a blend of semiochemicals released together, may have multiple functions, and, there-
fore, may be designated as a pheromone, allomone, kairomone, or synomone, depending on the context
in which it is being used or exploited.
15.3 Allomones: Defensive Chemicals ................................................................................................ 700
15.3.1 Range of Chemistry ...................................................................................................... 700
15.3.2 Effect of MTG and DAG Secretions against Natural Enemies .................................... 702
15.3.3 Other Behavioral Roles: Within Species ...................................................................... 702
15.4 Kairomones for Natural Enemies ................................................................................................ 703
15.4.1 Pentatomid Compounds as Kairomones for Egg Parasitoids ....................................... 704
15.4.2 Pentatomid Compounds as Kairomones for Diptera .................................................... 706
15.4.3 Pentatomid Compounds as Kairomones for Arthropod Predators ............................... 707
15.4.4 Predatory Stink Bugs Exploiting Semiochemicals of Their Prey ................................ 707
15.4.5 Tritrophic Interactions Involving Pentatomoid Bugs .................................................... 707
15.5 Overview: Semiochemicals in Life History of Pentatomoids, and Practical Applications ......... 708
15.5.1 Patterns of Production and Response to Pheromones .................................................. 708
15.5.1.1 Cross-Species Attraction ............................................................................... 709
15.5.1.2 Multiple Components: Ratios and Variability ................................................ 710
15.5.1.3 Dose and Release Rates of Stink Bug Semiochemicals ................................. 711
15.5.2 Current Role of Stink Bug Semiochemicals in Pest Management ................................ 712
15.5.3 Future Directions in Research and Applications ........................................................... 713
15.6 Acknowledgments ......................................................................................................................... 714
15.7 References Cited............................................................................................................................ 714