Brown marmorated stink bugs (Halyomorpha halys) are insects that are native to four Asian countries: China, Japan, Taiwan, and Korea. They have been introduced, however, in every country aside from Antarctica, Australia, and Africa. These stink bugs were accidentally introduced into Pennsylvania in 1996, where they then spread to almost the entirety of the United States and Canada within the next fifteen years. A half decade after, observations began occurring in nearly all European countries from their initial spread in France, Italy, and Switzerland. Russia and Chile were also invaded in the 2010's (Leskey and Nielsen, 2018). (Leskey and Nielsen, 2018)
Brown marmorated stink bugs are capable of living in a wide variety of habitats. Since their introduction to North America in the 1990's, they have been found in temperate areas as well as tropical ones, though less frequently (GBIF, 2021). They are unable to live in areas that are much too cold, and are uncommon in areas with an elevation over 1,200 meters (Holthouse et al., 2021). Additionally, brown marmorated stink bugs are often found in urban, suburban, and agricultural areas, where they are considered pests (GBIF, 2021). (GBIF, 2021; Holthouse, et al., 2021)
The adult brown marmorated stink bug is a medium-sized insect with mottled brown and tan coloration. They are differentiated from others in their family (Pentatomidae) by the black and white stripes on their abdomens and antennae. Males of the species are nearly identical to the females aside from a pair of claspers on the end of their abdomens that are used for mating. They have mouthparts specialized for sucking that are straw-like in appearance. Brown marmorated stink bugs experience five growth stages, or instars, before they reach adulthood. When they are at their smallest, the insects are black with reddish orange patterns on their backs, and are often mistaken for ticks. Over the course of the five instars, the red will fade to pale tan, and then to their namesake brown. Only adults of the species have fully functional wings, but developing wing pads will be visible in the fifth instar (Rice et al., 2014). (Holthouse, et al., 2021; Rice, et al., 2014)
These stink bugs experience hemimetabolous development, which is also known as incomplete development. They do not go through a pupal stage, but instead develop in various growth stages called instars. Brown marmorated stink bugs will develop with the same body plan and structure throughout their lifetimes. After hatching from their eggs, they will molt into larger versions of the same structure for five instars, slowly developing more adult-like features. They are considered adults in their final growth stage, after molting from the fifth instar. At this point, they will have fully grown and usable wings (Rice et al., 2014). (Rice, et al., 2014)
Brown marmorated stink bugs breed in the late spring to summer seasons, beginning in May and ending around August (Skvarla, 2017). During this time, a female will mate with multiple males, often consecutively. Males will do the same, but with nearly one hour breaks between each female he finds. Mating occurs most frequently late in the night and into early morning, and each insect can copulate nearly 20 times in one day. Females may be choosy with which male she selects, but only the males determine how long the breeding takes place for. As with most insects, brown marmorated stink bugs find mates through the use of pheromones that are most strongly released during the spring and summer months (Kawada and Kitamura, 1983). (Kawada and Kitamura, 1983; Skvarla, 2017)
Brown marmorated stink bugs have a fairly straightforward reproductive cycle that is reminiscent of other bugs in the family. They mate frequently during the breeding season that lasts from May to August, and will oviposit during this period as well (Skvarla, 2017). Females will lay around 454 eggs on average, give or take, during the season. These insects are sexual creatures, and males will fertilize the eggs internally following copulation. Some females will allow eggs to be fertilized multiple times before ovulation occurs (Kawada and Kitamura, 1983). (Kawada and Kitamura, 1983; Skvarla, 2017)
Brown marmorated stink bugs do not care for their young. The male is required only for fertilization of the eggs, and the female will simply gestate them, lay them, and leave (Kawada and Kitamura, 2017). (Kawada and Kitamura, 1983)
These insects spends most of their lives as adults as this stage can last an average of 5 months (Orkin, 2021). This is much longer than the approximately 50 days they will spend during their immature development (Kawada and Kitamura, 1983). The temperature will affect these numbers a great deal, however, as the bugs are incredibly sensitive to extreme hot and cold. Brown marmorated stink bugs will not make it to the adult stage if the temperature is outside of a range of 15°C to 30°C, with an optimum of 25°C. Development will be most efficient at this optimum, as well as more successful. Additionally, females will take longer to oviposit if the temperature is not suitable (Nielsen et al., 2008). (Kawada and Kitamura, 1983; Nielsen, et al., 2008; Orkin, 2021)
Brown marmorated stink bugs are not technically a social species, since they do not live together as families or have any sort of hierarchy between them. They will aggregate, however, at certain times in their lives or for specific reasons. For example, when the bugs are young, they may stay together with others in their brood for feeding, but will most certainly disperse eventually (GBIF, 2021). Alternatively, male brown marmorated stink bugs are capable of releasing and responding to an aggregation pheromone during any life stage. They will give off the pheremone typically in situations where they have found a resource that other stink bugs might also benefit from. This could be anything from an overwintering site to a food source. These insects are very sensitive to this pheromone, and will aggregate around the source within a radius spanning a few meters (Rice et al., 2014). (GBIF, 2021; Rice, et al., 2014)
Brown marmorated stink bugs are capable of not only extreme aggregation, but of long bouts of travel during both the day and night. The greatest distance flown in a single day by these bugs was 117 kilometers, but these cases are unusual. The average lies at around five or six kilometers in a day. Flights like these are very energetically costly, so they do not occur very frequently - perhaps only once or twice in a month. Nymphs of the species do not have fully developed wings, and therefore must travel by walking. Insects of the fifth instar have been recorded to walk as much as an impressive 20 miles in a span of four hours. Although it is intriguing that they can travel so far and so quickly, it can become problematic in agricultural and invasive settings (Leskey and Nielsen, 2018). (Leskey and Nielsen, 2018)
The brown marmorated stink bug's use of aggregation pheromones is crucial for the diapause behavior seen in this species in particular. They will aggregate at locations where they can keep warm over the winter while their bodies slow down and halt development in order to survive the chill. Brown marmorated stink bugs are very cold-sensitive, so places like hollowed trees, leaf-insulated soil, animal dens, and human-made structures make for perfect spots to hunker down for the cool seasons. One of the most important signals to begin diapause aside from temperature is day length, or photoperiod. When days begin to shorten, behaviors such as searching for and aggregation to locations such as those listed are onset (Leskey and Nielsen, 2018). (Leskey and Nielsen, 2018)
Brown marmorated stink bugs have no specific home range, as they are widely spread wherever they are found.
Brown marmorated stink bugs, like many other bugs in Hemiptera, communicate with vibrations and pheromones. The songs with which these insects communicate are created when their abdomens contact plants or other surfaces in a specific pattern. Another bug further down the surface will sense the vibration and be able to respond accordingly. Brown marmorated stink bugs prefer bendy plants such as beans because their vibrations are amplified by the flexibility, which is especially important in processes like looking for and impressing mates. Pheromone communication is also a big deal for brown marmorated stink bugs. As mentioned earlier, aggregation pheromones are an incredibly important tool that they use to find places at which to overwinter. When one insect finds a suitable location, it releases these pheromones to signal that it is adequate and others of the species should accumulate there (Mehal and Boyer, 2014). The antennae of brown marmorated stink bugs are invaluable for pheromone sensing as they possess olfactory sensilla, which are special cells that are designed to send electrical impulses to the insect's brain when they come in contact with a "smelly" molecule. This signal gives the bugs information about the chemical so that they can respond upon contact. Even more impressive is that, since the insects have two antennae, brown marmorated stink bugs are able to sense the strength and direction of the signal depending on the difference in how much of the chemical is picked up between each antenna (Saunders, 2019). (Mehal and Boyer, 2014; Saunders, 2019)
Another use of the antennae is as a tactile organ. Brown marmorated stink bugs can use them to assess the world around them, which is especially important for creatures that have limited sight capabilities. The bugs will also respond to being touched, and will fly away or move as necessary (Saunders, 2019). Additionally, the species is sensitive to various visual light cues. Day length, or photoperiod, is crucial for deciding when brown marmorated stink bugs should overwinter, as it gives them enough time to find a suitable site. It also allows them to know when to emerge from diapause and how to suit their circadian rhythm to the season (CABI, 2021). (CABI, 2021; Saunders, 2019)
Brown marmorated stink bugs are known to eat hundreds of different types of plants, making them very concerning insects (Leskey and Nielsen, 2018). They possesses piercing/sucking mouthparts in the shape of a long straw that is capable of drinking fluids possessed in plants and their fruits. Their mandibles have evolved to form stylets that surround the proboscis and support it, allowing it the strength to break into plant tissue (but thankfully not enough strength to harm an animal or a person) (Mehal and Boyer, 2014). (Leskey and Nielsen, 2018; Mehal and Boyer, 2014)
Most of the diet of brown marmorated stink bugs is comprised of arboreal plants (tree species), many of which are not natural hosts of these insects. Some of the most important of these, especially for the bugs' rapid spread, include species within the Acer genus (Ash trees) and the Fraxinus genus (Maples), as well as the foxglove/empress tree (P. tomentosa), and the varnish tree (A. altissima). In addition, brown marmorated stink bugs are important agricultural pests, preying on victims such as peaches, pears, wheat, grapes, apples, soybean, sorghum, corn, tomatoes, okra, peppers, and eggplants. Typically, these insects prefer to feed on the fruit of their host plant, but they have been known to also ingest sap from the leaves and stems (Rice et al., 2014). (Rice, et al., 2014)
Brown marmorated stink bugs have little need for anti-predator adaptations outside of their native range since they are a largely uncontrolled species without natural enemies. However, their brown mottled coloration may assist in camouflaging the bug in leaf litter and on tree bark (Holthouse et al., 2017). Brown marmorated stink bugs are known to have many predators from birds to slugs; rats to other insects. Various generalist birds and small carnivorous mammals will snack on the small arthropods, but their biggest predators are typically other insects and spiders (Gonzales et al., 2017). Jumping spiders (family Salticidae are the most well known of the arachnid predators, but certainly not the only ones. Insects such as grasshoppers, katydids, lacewings, soldier beetles, and even other species of stink bugs will predate on brown marmorated stink bugs. A large portion of enemies are wasps, however, since there are quite a few species of parasitoids that will oviposit inside the eggs of brown marmorated stink bugs. They will hatch, develop, and pupate all within the egg, making the insect crucial to their survival (Schoof, 2020). Arguably the most important of the natural enemies is the samurai wasp (Trissolcus japonicus). This egg parasitoid greatly limits egg development and is often considered as an option for biological control (Holthouse et al., 2017). Other species of parasitoids that affect the life of brown marmorated stink bugs are also in the Trissolcus species (T. edessae, T. eushisti, T. brochymenae, T. itoi, T. mitsukurii, T. plautiae). Additionally, some chalcid wasps (Anastatus mirabilis, Anastatus ruduvii) parasitize the bugs as well (Schoof, 2020). (Gonzales, et al., 2017; Holthouse, et al., 2017; Schoof, 2020)
Although brown marmorated stink bugs are mainly viewed as a pest species, they do have a few roles in ecosystems beyond eating/damaging plants and fruit. Firstly, these insects are a food source for a variety of other animals including birds, rats, and other insects (see above list) (Gonzales et al., 2017). Also, the insect's eggs are an important host for various parasitoid predators as listed below (Schoof, 2020). (Gonzales, et al., 2017; Schoof, 2020)
As with many other insect species, gut symbionts are important in digestive function for brown marmorated stink bugs. The organism will aid the stink bugs in digesting their food, and in turn will benefit with a viable host as a home. Of these symbionts is one species in particular: P. agglomerans. This bacterium creates a gut film for the insects that assists in protection for the gut lining and therefore better digestion. Brown marmorated stink bugs acquire this symbiont by ingesting them off of their eggs after they hatch. This is because gut symbionts rub off in a residue from the mother's anus as she lays her eggs (Taylor et al., 2014). Another symbiont important to brown marmorated stink bugs is P. carbekii, a microbe whose presence correlates to the insects having longer lifespans with shorter development times, as well as more regular behavior and better reproductive potential (Taylor, 2016). Another bacterium that lives within the brown marmorated stink bugs is a lot more dangerous. Paulownia Witches' Broom (PaWB) is a pathogen vectored (transmitted, alternatively) by the insects that ruins the bark of the empress tree (Paulownia tomentosa, among others), furthering the bugs' pest status (Mitchell, 2004). (Mitchell, 2004; Taylor, 2016; Taylor, et al., 2014)
Perhaps the only benefit brown marmorated stink bugs provide for humans is their potential as research subjects, especially in the area of control and natural enemy ecology. The sheer amount of damage they are capable of makes their study crucial within the agricultural and entomological community, thus creating an economical niche (CABI, 2021). (CABI, 2021)
Thankfully, brown marmorated stink bugs cannot physically "bite" humans as their mouthparts are not strong enough to break through skin (Mehal and Boyer, 2014). However, allergic reactions have been recorded to occur as a result of the defensive chemicals the bugs produce and dermatitis is possible if the insect's hemolymph gets on the skin (Skvarla, 2017). (Mehal and Boyer, 2014; Skvarla, 2017)
Brown marmorated stink bugs are pests of plant species. Large sums of money are spent yearly to attempt to control these insects due to their status as invasive and damaging in many parts of the world. They cause severe damage to agricultural crops, ornamentals, and native species alike. There would be an over 40 billion dollar loss in the United States alone if brown marmorated stink bugs attacked all vulnerable crops. Secondary pest outbreaks and food contamination are a side affect of invasion as well, especially since brown marmorated stink bugs can vector multiple pathogens (CABI, 2021). One of the biggest visible impacts is damage to the fruit from both feeding damage and bacteria, which make it unmarketable or inedible (Rice et al., 2014) Even worse is the bugs' apparent growing resistance to some market pesticides (Suits, 2015). A large increase in pesticide use is partly to blame for this, as brown marmorated stink bugs are excellent at flying, climbing, and avoiding residues, thus requiring larger coverage. These insects are a problem in their native range in Asia as well, but have more natural enemies to keep the species in check there (Holthouse et al., 2017). Additionally, being invasive with no natural enemies present in their non-native range, brown marmorated stink bugs are a massive ecological threat as well. They damage native species and outcompete other native insects and arthropods for food sources and shelter. Unfortunately, it is a long and expensive process with no clear outcome to introduce a biological control, and brown marmorated stink bugs proliferate too quickly for mechanical controls to make much of a dent in their numbers (Servick, 2018). (CABI, 2021; Holthouse, et al., 2017; Servick, 2018; Suits, 2015)
Lastly, brown marmorated stink bugs are considered household pests. Aside from feeding on backyard gardens and window-box plants, these insects are nuisances in that they will move into homes for the winter, often in massive quantities. Since they require a warm place to undergo diapause, a human dwelling is ideal. Therefore, especially in rural settings, these bugs will be difficult to evacuate and often require costly pest control. Thankfully, they do not do much damage aside from causing a lingering odor (CABI, 2021). (CABI, 2021)
Brown marmorated stink bugs are common where they are found, so they have no special conservation status.
Amy Bagby (author), Colorado State University, Amy Bagby (editor), Colorado State University.
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
living in the southern part of the New World. In other words, Central and South America.
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
living in landscapes dominated by human agriculture.
Referring to an animal that lives in trees; tree-climbing.
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
uses smells or other chemicals to communicate
active at dawn and dusk
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.
union of egg and spermatozoan
an animal that mainly eats leaves.
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
an animal that mainly eats fruit
An animal that eats mainly plants or parts of plants.
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.
Found in northern North America and northern Europe or Asia.
fertilization takes place within the female's body
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
having the capacity to move from one place to another.
the area in which the animal is naturally found, the region in which it is endemic.
active during the night
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
chemicals released into air or water that are detected by and responded to by other animals of the same species
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
breeding is confined to a particular season
reproduction that includes combining the genetic contribution of two individuals, a male and a female
living in residential areas on the outskirts of large cities or towns.
uses touch to communicate
Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
Living on the ground.
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
living in cities and large towns, landscapes dominated by human structures and activity.
movements of a hard surface that are produced by animals as signals to others
uses sight to communicate
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