Invasive Species Compendium

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Datasheet

Dendroctonus frontalis
(southern pine beetle)

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Datasheet

Dendroctonus frontalis (southern pine beetle)

Summary

  • Last modified
  • 28 March 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Dendroctonus frontalis
  • Preferred Common Name
  • southern pine beetle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Field symptoms of an infestation by D. frontalis (large southern pine beetle) at Little Lake Creek Wilderness Area, Sam Houston National Forest, south-east Texas, USA.
TitleField symptoms
CaptionField symptoms of an infestation by D. frontalis (large southern pine beetle) at Little Lake Creek Wilderness Area, Sam Houston National Forest, south-east Texas, USA.
CopyrightRobert N. Coulson/Texas A&M University
Field symptoms of an infestation by D. frontalis (large southern pine beetle) at Little Lake Creek Wilderness Area, Sam Houston National Forest, south-east Texas, USA.
Field symptomsField symptoms of an infestation by D. frontalis (large southern pine beetle) at Little Lake Creek Wilderness Area, Sam Houston National Forest, south-east Texas, USA.Robert N. Coulson/Texas A&M University

Identity

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Preferred Scientific Name

  • Dendroctonus frontalis Zimmermann, 1868

Preferred Common Name

  • southern pine beetle

Other Scientific Names

  • Dendroctonus arizonicus Hopkins, 1909

International Common Names

  • English: beetle, southern pine
  • French: dendroctone méridional du pin

EPPO code

  • DENCFR (Dendroctonus frontalis)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Scolytidae
  •                             Genus: Dendroctonus
  •                                 Species: Dendroctonus frontalis

Notes on Taxonomy and Nomenclature

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Dendroctonus frontalis Zimmermann was originally described by Zimmermann in 1868. In 1963, Stephen Wood synonymized D. arizonicus Hopkins and D. mexicanus Hopkins with D. frontalis Zimmermann.

Vité et al. (1974) provided conclusive evidence, that D. frontalis and D. mexicanus are two separate species. Their findings prompted Wood (1974) to reinstate D. mexicanus as a valid species.

The systematic history of the species was recently reviewed by Cognato (2011), who provided a detailed overview of D. frontalis from morphological, biological, karyological and molecular data.

Armendariz-Toledano et al. (2014) found that two newly discovered apparent morphological variants of D. frontalis from Central America and southern Mexico may actually represent two distinct sibling species. 

Description

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D. frontalis is a holometabolous insect (see Pictures). The life stages include egg, four larval instars, pupa, and adult. Basic facts about the morphology of these stages follow, from Payne (1980).

The egg is slightly oblong to oval with rounded ends. It is ca. 1.5 mm long and 1 mm wide, opaque, pearly white, and shiny.

The larva is a subcylindrical, wrinkled, legless grub with three thoracic and 10 abdominal segments. It is yellowish white in colour. Early stage larva are ca. 2 mm in length and mature larvae ca. 5 to 7mm. The mouth parts are well developed with stout and dark-coloured mandibles.

The exarate pupa is yellowish white in colour and is 3 to 4 mm in length. It has the general form of the adult but with wing pads and legs folded beneath and the abdominal segments exposed.

The adult is cylindrical and stout to elongated in shape. It is 2 to 4 mm in length and brownish to black in colour. The front of the head is coarsely punctured and channelled in both sexes. The adults have clubbed antennae, typical of Scolytidae, and the elytral declivity is convex. Females are distinguished from males by the presence of a mycangium on the anterior pronotum. Males lack the mycangium but have a distinct frontal groove. Also, the elevations or tubercles on the male head are more distinct than in females.

Distribution

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Historically, D. frontalis has been distributed primarily in the pine forest regions of the southern USA (Alabama, Arkansas, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, Oklahoma, South Carolina, Tennessee, Texas, and Virginia). In Central America: Belize, El Salvador, Guatemala, Honduras, Nicaragua. However, the outbreak area for D. frontalis is expected to expand into more northern latitudes as a consequence of projected global warming trends (Williams and Liebhold, 2002).

Also see CABI/EPPO (2002).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

IsraelAbsent, formerly presentCABI/EPPO, 2002; EPPO, 2014

North America

MexicoRestricted distributionNativeCABI/EPPO, 2002; EPPO, 2014
USARestricted distributionNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-AlabamaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-ArizonaPresentCABI/EPPO, 2002; EPPO, 2014
-ArkansasPresentNativeCABI/EPPO, 2002; EPPO, 2014
-CaliforniaPresentCABI/EPPO, 2002
-DelawarePresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-District of ColumbiaPresentCABI/EPPO, 2002; EPPO, 2014
-FloridaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-GeorgiaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-KentuckyPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-LouisianaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-MarylandPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-MississippiPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-North CarolinaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-OklahomaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-PennsylvaniaPresentCABI/EPPO, 2002; EPPO, 2014
-South CarolinaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-TennesseePresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-TexasPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-VirginiaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
-West VirginiaPresentPayne, 1980; CABI/EPPO, 2002; EPPO, 2014

Central America and Caribbean

BelizePresentNativeCABI/EPPO, 2002; EPPO, 2014
El SalvadorPresentNativeCABI/EPPO, 200202; EPPO, 2014
GuatemalaPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
HondurasPresentNativePayne, 1980; CABI/EPPO, 2002; EPPO, 2014
NicaraguaPresentNativeCABI/EPPO, 2002; EPPO, 2014

Europe

IrelandAbsent, invalid recordCABI/EPPO, 2002; EPPO, 2014

Hosts/Species Affected

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The hosts of D. frontalis include a variety of Pinus species. In the southern USA the prominent hosts include the commercially important species of southern yellow pine: Pinus echinata, P. taeda, P. palustris, and P. elliotti. However, the insect also infests: P. rigida, P. virginana, P. pungens, P. strobus, P. glabra, P. serotina. In Central America important hosts include: P. caribaea, P. englemannii, P. leiophylla, P. maximinoi, and P. oocarpa. It is worthy noting that in outbreak conditions, D. frontalis will utilize a variety of coniferous hosts including exotic species of Pinus as well as Picea species and Tsuga species. Among the southern yellow pines, longleaf pine, P. palustris, is highly resistant to colonization by D. frontalis.

Growth Stages

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Symptoms

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Herbivory by D. frontalis results in mortality to host pines and the foliage changes in colour from green to bright red as the trees desiccate. Infestations can be detected by aerial survey, which is the most common way that outbreaks are monitored.

Winding S-shaped egg galleries can be seen in the cambium layer.

Pitch tubes, about the size and colour of popcorn can be seen along the entire length of the trunk. Pitch tubes may not be present in very weak trees, but reddish boring dust in bark crevices can indicate infestation (Swain and Remion, 1981)
 

List of Symptoms/Signs

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SignLife StagesType
Leaves / yellowed or dead
Stems / gummosis or resinosis
Stems / internal feeding
Whole plant / plant dead; dieback

Biology and Ecology

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Natural History

Coulson (1980) and Flamm et al. (1988) summarized the significant features of the life cycle through which D. frontalis passes, with particular references to strategies influencing survival, growth and reproduction. The natural history can be partitioned to include activities associated with host selection, aggregation and attack, mating and oviposition, reemergence, brood development, and emergence.

Host selection is initiated by female beetles. The behaviour that guides host selection is not fully understood. Response to both olfactory and/or visual cues has been proposed as the mechanism. Once a few beetles have bored into a susceptible host, secondary attraction takes place. This aggregation and attack phase is a critical juncture in the natural history of D. frontalis, as it serves to concentrate dispersing beetles in numbers large enough and in a time frame short enough to overcome the hosts defence mechanisms. In order for a brood population to survive the host tree must be killed.

Pinus species have three defense systems that serve to deter colonisation by D. frontalis: wound-cleansing, performed by the resin duct system; containment of infection, performed by hypersensitive reaction; and wound-healing through the formation of periderm and callus tissue. The first line of defense is the resin system and both the physical and chemical properties of the resin and the flow rate interact to discourage colonisation. During the aggregation phase, D. frontalis also introduces symbiotic fungi and other microorganisms. The microorganisms introduced by the beetle are thought to be important in both overcoming host resistance and also in larval nutrition.

Aggregation behavior is guided by a suite of beetle and host-produced compounds that act initially to attract beetles to the host and later to inhibit response to the tree. A variety of natural enemies and associated organisms also respond to the attractants. The process of attack is initiated by females who are joined by males. The duration of attack can range from 8 days to 6 weeks.

Mating and oviposition take place within the successfully attacked host tree. Females initiate excavation of egg galleries in the phloem. Copulation takes place within a nuptial chamber. D. frontalis is monogamous within an individual host tree. After mating, the female excavates a characteristic S-shaped egg gallery (see Pictures). Eggs are laid in niches at precise intervals along the lateral walls of the gallery. The insect regulates egg population size and thereby ensures sufficient food resources for developing larvae. Once oviposition is completed, adult beetles either die in the gallery or re-emerge. Re-emergence behaviour is significant in that females are capable of establishing multiple cohorts.

Brood development (from egg to adult) ranges from ca. 26 to 140 days depending on temperature. Eggs usually hatch in 3 to 27 days. Larvae excavate galleries perpendicular to the egg gallery. There are four larval stages. The first three forage in the phloem region and the fourth moves into the outer corky bark. Larval development is completed in 13 to 63 days. Larvae pupate in the outer bark and later transform into callow adults in 3 to 36 days. Mature adults emerge through exit holes bored from the pupal chamber. If weather conditions are unfavorable the adults will remain in the bark. The final phase in the life cycle is emergence, which occurs in small daily increments over a 14 to 28 day period. Beetle dispersal takes place in the morning and afternoon hours.

Depending on weather conditions, D. frontalis is capable of producing 6 to 8 generations per year. Explosive changes in population size can occur as a consequence of the short duration of the life cycle and it is for this reason that D. frontalis is considered among the most damaging of forest insect pests.

Population growth in infestations

Populations of D. frontalis occur in clumps of infested hosts (infestations) distributed throughout a forest. Typically, adult beetles emerging from an infested tree attack the nearest neighbouring host. This behaviour results in the development of multiple-tree infestations, which are often referred to as spots (see Pictures). Generally, an infestation is comprised of a number of trees containing beetles in different stages of development. Infestations of D. frontalis can be distinguished from the surrounding vegetation, as the crowns of trees become discoloured following colonisation by the insects.

Herbivory by D. frontalis kills the host pines and the foliage changes in colour from green to bright red as the trees desiccate. Infestations are commonly detected and monitored by aerial survey. Individual infestations are often observed to grow in size as new hosts are colonised. Individual infestations are usually established in the early spring and growth takes place through the summer and continues into the fall. This pattern of continuous infestation growth is a unique attribute of D. frontalis population ecology. Under epizootic conditions, individual infestations often coalesce and the pattern and extent of tree mortality substantially modify the structure of the forest landscape (Coulson and Wunneburger, 2000; Perkins and Matlack, 2002) (see Pictures). In most instances, infestations cease to expand towards the end of the summer and adult populations disperse. The annual number of infestations observed for a geographic region varies by orders of magnitude. In eastern Texas, where survey records have been maintained since 1960, the number of infestations recorded in a given year has ranged from none to more than 17,000 (Coulson et al., 1999).

Interaction with other bark beetle species

D. frontalis is a member of a guild of bark beetles that includes D. terebrans (Oliver), (the black turpentine beetle), Ips avulsus (Eichhoff) (the four-spined engraver), I. grandicollis (Eichhoff) (the eastern five-spined engraver), and I. calligraphus (Germar) (the six-spined engraver). It is common to find all members of the guild in an infested tree. Generally, the individual guild members partition the host to avoid competition for food and habitat resources. Under enzootic conditions behaviour associated with resource partitioning among the guild members is relaxed and it is common to observe intermingling of the various species along the infested bole of the host tree. Although each guild member is capable of infesting hosts in the absence of D. frontalis, large outbreaks are uncommon. Rather, they colonise single trees or small groups of trees, generally in association with lightning strikes, logging, windthrow, fire, naval stores (resin tapping), or other human-caused disturbances.

Epizootiology

Forest stand and site conditions play an important role in the distribution and abundance of D. frontalis infestations. Several different methods have been devised to classify forest stands according to the hazard posed by D. frontalis. Each rating system has novel features, but all involve integration of a subset of variables: tree species, radial growth, height, DBH, stand basal area (pine, hardwood, total), species composition, site index, degree of crown closure, and landform classification. In general, stands growing on poor sites and containing a mature susceptible host species (e.g., P. echinata or P. taeda in the southern US) with high basal area and stagnant radial growth are considered to be high hazard. These stands represent habitat patches and they are suitable for D. frontalis infestation growth. Most of the annual timber volume loss attributed to the insect occurs in high hazard stands. The high hazard habitat patches are often spatially separated by forest conditions that do not support D. frontalis survival, growth, and reproduction. The habitat patches are linked via the dispersal behaviour of adult insects. In instances where populations are at enzootic levels or habitat patches occur at distances beyond the dispersal capacity of the insect, lighting-struck hosts can serve as refuges and stepping stones (Lovelady et al., 1991). When a lightning-struck host occurs in a habitat patch it can serve as an epicenter for an infestation. Lightning-struck hosts are an important component of the natural history of D. frontalis. These hosts have greatly diminished defence capacity and can be successfully colonised by populations too small to overcome a healthy tree. It is thought that the insect responds to resin odours associated with the wound produced by lightning striking the tree. The lightning-struck hosts are also recognised and colonised by the other members of the bark beetle guild. Other types of disturbed hosts can also function as refuges, stepping stones, and epicentres for infestations (Coulson et al., 1999).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Alcaligenes faecalis Pathogen
Aspergillus flavus Antagonist
Beauveria bassiana Pathogen
Coeloides pissodis Parasite
Corticeus glaber Predator
Corticeus parallelus Predator
Dendrosoter sulcatus Parasite
Diathrombium diaphane Parasite
Dinotiscus dendroctoni Parasite
Dryocopus pileatus Predator
Enterobacter aerogenes Pathogen
Eurytoma Parasite Larvae/Pupae
Haematonectria haematococca Pathogen
Heydenia unica Parasite
Lyctocoris elongatus Predator
Medetera bistriata Predator
Megophthrombium gracile Parasite
Metarhizium anisopliae Pathogen
Paecilomyces viridis Pathogen
Picoides pubescens Predator
Picoides villosus Predator
Platycryptus undatus Predator USA; Texas Pinus
Pyemotes dryas Parasite
Pyemotes giganticus Parasite
Rhopalicus pulchripennis Parasite
Roptrocerus xylophagorum Parasite
Scoloposcelis flavicornis Predator
Serratia marcescens Pathogen
Spathius pallidus Parasite
Thanasimus dubius Predator
Thanasimus formicarius Predator West Virginia Pinus
Unikaryon minutum Pathogen
Verrucosa arenata Predator USA; Texas Pinus

Notes on Natural Enemies

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D. frontalis has a suite of natural enemies and associates. The natural enemies primarily come from Coleoptera, Hymenoptera, and Diptera. Within Coleoptera there are important predators in the families Trogositidae (Temnochila and Tenebroides) and Cleridae (Thanasimus). Predation by Thanasimus dubius is particularly noteworthy, and both adults and larvae are predators. Survey entomologists often observe this insect in great numbers on trees being colonised by D. frontalis. There are several families of hymenopterous parasitoids associated with D. frontalis. Most occur within the families Braconidae (Cenocoelius, Coeloides, Atanycolus, Dendrosoter, Meteorus, Spathius), Pteromalidae (Dinotiscus, Roptrocerus, Heydenia), Eurytomidae (Eurytoma), and Platygastridae (Platygaster, Leptacis). Prominent Diptera include representatives from the Stratiomyiidae (Zabrachia), Dolichopodidae (Medetera), Sciaridae, and Lonchaeidae (Lonchaea). These species, together with various microbes and predacious mites, undoubtedly provide some degree of D. frontalis population regulation. There continues to be a great deal of research interest in developing biological control tactics for D. frontalis.

Plant Trade

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Plant parts not known to carry the pest in trade/transport
Bark

Impact

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The direct economic impact of the insect includes loss of revenue associated with infested trees as well as the cost of suppression tactics. In commercial forests, management plans and schedules are also disrupted. As the insect infests and kills mature trees, the economic loss occurs when the stands are at their most valuable. During outbreaks the markets for infested trees become saturated. The value of infested host material declines to a point where it is often not profitable to salvage and trees are left to decompose in the forest. This situation has commonly occurred in the large outbreaks reported in Mexico and Central America. In addition, the infestations may occur in remote sites and therefore be inaccessible to salvage crews. As D. frontalis is typically found in areas where climatic conditions are mild, the rate of decomposition of infested trees is rapid, so there is only a brief period when infested trees are merchantable. Furthermore, the blue-staining fungi, introduced by D. frontalis, discolour the wood and make it undesirable as a source of pulp for certain types of paper products.

Environmental Impact

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The principal ecological impact of D. frontalis is as a disturbance agent that triggers ecological succession in old-age pine forests. In these forests net primary production has stagnated, nutrients are concentrated and bound, and plant and animal diversity is low. In natural forests, infestations resulted in disturbance patches that led to release and reorganization of the forest landscape. The nutrients bound as pine biomass were liberated through decomposition processes initiated by D. frontalis herbivory. The open patches became sinks for early successional plant and animal species and biodiversity was re-introduced into the forest landscape. The patches also provided both habitat and food resources for a variety of wildlife species. The infestations introduced age-class diversity and thereby influenced the kind, size, shape, and number of vegetation patches that formed the forest landscape mosaic (Coulson et al., 2000).

Social Impact

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The social impacts of D. frontalis occur when the insect infests trees in urban and suburban settings and when it influences recreational use of forest landscapes. In urban and suburban environments, remnant pine forests are often used for housing developments. Old-growth pines are particularly valued on home sites for their aesthetic beauty and shade. Building construction practices that alter drainage pathways, compact soil, damage roots and injure standing pine trees often result in the creation of D. frontalis infestations in urban settings. Aside from the emotional distress created by the loss of the trees, property values are often greatly affected as well. In forests, recreationists will often avoid campsites where D. frontalis infestations are present. Snags, resulting from infested trees left in the forest, are dangerous to campers, hikers, and foresters.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

There are four methods used in direct suppression of southern pine beetle: salvage removal, cut-and-leave, pile-and-burn, and chemical insecticides (Swain and Remion, 1981). Accessibility to the infestation, value of infested host material, markets, and management constraints associated with forest ownership and management objectives generally dictate which procedure is used. In addition to direct control procedures, another approach to reducing the impact of D. frontalis is to avoid the conditions that promote infestations. There are a number of steps that can prevent outbreaks of the insect.

Salvage removal involves cutting and marketing infested trees. Typically a buffer of green uninfested trees surrounding the infestation is included in the salvage. The rational for the buffer strip is to ensure that newly infested trees, that could serve to fuel the establishment of a new infestation, are not overlooked. Forest managers prefer salvage removal as it provides some revenue for the infested timber. However, this approach is often impractical because: infestations may not be accessible, the volume of timber may be insufficient, the quality of wood may have deteriorated, there may not be a market for infested wood (a typical situation during outbreaks), or the infestation may occur in a sensitive area (e.g., a wilderness area or a National Forest). Salvage removal may also take longer to implement than the other methods.

Cut-and-leave is a suppression tactic that involves felling infested trees and a buffer of uninfested trees and leaving them in the forest. This procedure is recommended for small infestations (in the range of 10 to 50 trees) that are actively growing (expanding), for example, during months of May to October in the southern US. Although the mode of action of this approach is the subject of some speculation, the treatment is thought to disrupt infestation growth and result in dispersal of adults into the surrounding environment. Empirical research has indicated that new infestations do not appear around cut-and-leave treatments, in numbers greater than would be expected without treatment (Fitzgerald et al., 1994).

Cut-pile and burn is a suppression tactic that involves felling infested trees, gathering them into a pile, and burning them. This procedure destroys the insects in situ. This procedure is rarely used because it is expensive to implement, there may be environmental constraints associated with air pollution, and there is a risk of forest fire. This procedure is used sometimes as a component of site regeneration.

Chemical control is a suppression tactic that involves applying a registered pesticide to trees infested by the insect. This tactic is useful for small infestations or individual infested trees that occur on high-value sites (e.g., in urban forests, recreational areas, parks, etc.). This tactic requires special application equipment and the insecticides are expensive. In the USA, the registered insecticides approved for suppression of D. frontalis are limited and subject to change. The USDA Forest Service, Forest Health Protection, Cooperative Extension, or the state forestry agencies can provide information on approved insecticides.

Prevention of D. frontalis outbreaks involves application of forest management procedures that create conditions unsuitable for the establishment and growth of infestations. As mentioned above, most of the economic loss caused by the insect occurs in high hazard stands, i.e., stands stocked with a preferred host tree species, 40 years of age or older, growing on poor sites, with high basal area, and stagnant growth. These conditions can be modified by selection of less vulnerable host species, e.g., Pinus palustris (longleaf pine); site preparation practices that favour vigorous growth, e.g., fertilisation, open spacing, proper soil drainage; thinning to reduce basal area; and short rotation harvesting (e.g. 35 to 40 years).

References

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Armendáriz-Toledano F, Niño A, Sullivan BT, Macías-Sámano J, Víctor J, Clarke SR, Zúñiga G, 2014. Two species within Dendroctonus frontalis (Coleoptera: Curculionidae): evidence from morphological, karyological, molecular, and crossing studies. Annals of the Entomological Society of America, 107(1):11-27. http://esa.publisher.ingentaconnect.com/content/esa/aesa/2014/00000107/00000001/art00002

CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp.

CABI/EPPO, 2002. Dendroctonus frontalis. Distribution Maps of Plant Pests, No. 634. Wallingford, UK: CAB International.

Cognato AI, 2011. A review of Dendroctonus frontalis Zimmermann systematics. General Technical Report - Southern Research Station, USDA Forest Service, No.SRS-140:7-12.

Coulson RN, 1980. Population dynamics of the southern pine beetle. In: Thatcher RC, Searcy JL, Coster JE, Hertel GD, eds. The Southern Pine Beetle. USDA For. Serv. and Sci. and Educ. Admin. Tech. Bull., 1631, 71-105.

Coulson RN, McFadden BA, Pulley PE, Lovelady CN, Fitzgerald JW, Jack SB, 1999. Heterogeneity of forest landscapes and the distribution and abundance of the southern pine beetle. Forest Ecology and Management, 114(2/3):471-485; 2 pp. of ref.

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Fitzgerald JW, Couslon RN, Pulley PE, Flamm RO, Oliveria FL, Swain KM, Drummond DB, 1994. Suppression tactics for Dendroctonus frontalis Zimmerman (Coleoptera: Scolytidae): an examination of the occurrence of infestations adjacent to treatment sites. Journal of Economic Entomology, 87(2):417-425

Flamm RO, Coulson RN, Payne TL, 1988. The southern pine beetle. In: Berryman AA, ed. Dynamics of Forest Insect Populations. Plenum, 531-553.

Klepzig KD, Moser JC, Lombardero MJ, Ayres MP, Hofstetter RW, Walkinshaw CJ, 2001. Mutualism and antagonism: ecological interactions among bark beetles, mites and fungi. Biotic interactions in plant-pathogen associations, 237-267; 69 ref.

LeConte JL, 1876. Appendix to Zimmermann's synopsis of the Scolytidae of American north of Mexico. Trans-Amer. Entomol. Soc. 2:150-178.

Lovelady CN, Pulley PE, Coulson RN, Flamm RO, 1991. Relation of lightning to herbivory by the southern pine bark beetle guild (Coleoptera: Scolytidae). Environmental Entomology, 20(5):1279-1284

MNREI, 2000. Pine trees under attack. The Ministry of Natural Resources, the Environment and Industry. Government of Belize. BioFocus, 2:10.

Payne TL, 1980. Life history and habits. In: Thatcher RC, Searcy JL, JE Coster, Hertel G, eds. The Southern Pine Beetle. USDA For. Serv. and Sci. and Educ. Admin. Tech. Bull., 1631, 7-28.

Perkins TE, Matlack GR, 2002. Human-generated pattern in commercial forests of southern Mississippi and consequences for the spread of pests and pathogens. Forest Ecology and Management, 157(1/3):143-154; many ref.

Swain KM, Remion MC, 1981. Direct control methods for the southern pine beetle. USDA Comb. For. Pest Res. and Dev. Prog. Agric. Handb. No. 575, 15.

Vite JP, Islas SF, Renwick JAA, Hughes PR, Kliefoth RA, 1974. Biochemical and biological variation of southern pine beetle populations in North and Central America. Zeitschrift fur Angewandte Entomologie, 75(4):422-435

Williams DW, Liebhold AM, 2002. Climate change and the outbreak ranges of two North American bark beetles. Agricultural and Forest Entomology, 4(2):87-99; 49 ref.

Wood SL, 1963. A revision of the bark beetle genus Dendroctonus Erichson (Coleoptera: Scolytidae). The Great Basin Naturalist, 23:1-117.

Wood SL, 1974. New synonymy and records of American bark beetles (Coleoptera: Scolytidae). Great Basin Naturalist, 34(4):277-290

Zimmermann C, 1868. Synopsis of the Scolytidae of America north of Mexico, with notes and an appendix by J.L. LeConte, Am. Entomol. Soc. Trans. 2:149.

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