Ips grandicollis
(five-spined bark beetle)

Pictures

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Identity

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Preferred Scientific Name

• Ips grandicollis (Eichhoff, 1868)

Preferred Common Name

• five-spined bark beetle

Other Scientific Names

• Ips cacographus (LeConte, 1868)
• Ips chagnoni Swaine, 1916
• Ips cloudcrofti Swaine, 1924
• Ips cribricollis (Eichhoff)
• Tomicus cribricollis Eichhoff, 1869
• Tomicus grandicollis Eichhoff, 1868

International Common Names

• English: bark beetle, five- spined; engraver, southern pine; southern pine engraver
• French: scolyte à grand corselet

Local Common Names

• USA: five spined pine engraver

Taxonomic Tree

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

Description

Top of page Eggs

Eggs are 0.7-0.8 mm long, 1.5 times as long as wide; smooth, oval and translucent white.

Larvae

Larvae are white, legless grubs with highly sclerotized heads. Larvae tend to assume the shape of the letter 'C'. The larvae develop through three instars, which are identical except for size.

Pupae

White pupae are found in frass-encircled chambers at the end of the larval galleries, where they remain responsive and in constant motion. Pupae display many rudimentary adult features, such as legs, elytra and antennae.

Adults

Adults are reddish-brown to dark-brown, 2.9-4.6 mm long and 2.7 times as long as wide. The body is cylindrical with rounded lateral margins; the posterior margin has a deeply excavated declivity with elevated margins; the lateral margin is armed with spines, five on each elytron. When viewed from above, the head is hidden by the pronotum. Antennae have a five-segmented funicle and a flat, imperfectly circular club with two visible sutures.

Distribution

Top of page I. grandicollis is known from the USA and Canada since 1868. As well as those states listed (see Distribution List), I. grandicollis is probably present in Delaware, Kentucky and Vermont because these states contain suitable hosts and are surrounded by states where I. grandicollis is known to occur.

More recently discovered in more southerly locations, such as Central America, Caribbean Islands and Mexico (Garraway, 1986; Berrios et al., 1987; Guerra et al., 1989; Haack and Paiz-Schwartz, 1997) but it is not known whether I. grandicollis is native to these regions or introduced. I. grandicollis was introduced to Australia in the 1940s, where it is now widespread and common throughout forests that contain Pinus (Rimes, 1959; Morgan and Griffith, 1989; Abbott, 1993).

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.

Risk of Introduction

Top of page I. grandicollis is an A1 quarantine pest for EPPO. As I. grandicollis can make primary attacks on Pinus spp., it does present a risk to the EPPO region, where Pinus spp. are important forest trees. This risk is regarded as relatively moderate. However, following its introduction into Australia, I. grandicollis has damaged Pinus pinaster and P. radiata, which are also widely planted in the EPPO region (EPPO/CABI, 1997).

Indigenous Ips spp. already occur on conifers throughout most of the EPPO region, so the risk arising from introduced species is uncertain (EPPO/CABI, 1997).

Hosts/Species Affected

Top of page I. grandicollis is found beneath the bark of dead and dying trees, feeding and reproducing in the phloem of nearly all species of Pinus within its geographical range.

Host Plants and Other Plants Affected

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Growth Stages

Top of page Post-harvest, Vegetative growing stage

Symptoms

Top of page The first signs of attack are multiple small accumulations of fine, light-coloured frass upon the surface and in crevices in the bark of the bole, where it collects after being excavated from burrows drilled into the phloem by attacking adults. The burrows themselves are often not visible without removing bark flakes. Trees develop a 'red flag' appearance in the crown, as needles in branches above attack sites begin to turn red. In the early and mid stages of attacks, nuptial chambers and brood galleries with adults and their developing progeny can be observed if bark is peeled to expose the phloem near frass accumulations. Later on, the entire crown turns red and then brown. The bark becomes riddled with 1.5-mm emergence holes made by emerging offspring and can be pulled off readily due to beetles and other insects mining the phloem. By this stage, most I. grandicollis will be gone.

Once under attack, a tree very rarely survives, although it is not always easy to establish whether I. grandicollis actually killed the tree, rather than other co-attacking species, or whether the tree was dying from other causes that predisposed it to bark beetle attack.

List of Symptoms/Signs

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Biology and Ecology

Top of page Adult males locate potential host material through a process that is not well understood. Nearly all recently dead host material gets colonized; lightning strikes and windthrown trees are usually found and colonized, as is most logging debris. However, living and apparently healthy trees can also be attacked. Unlike most of its congeners, I. grandicollis is clearly attracted to host volatiles, which presumably contributes to host location. Once a tree has been chosen, the male bores through the bark and excavates a nuptial chamber in the phloem layer, while emitting in his frass the aggregation pheromone ipsenol (Renwick and Vite, 1972). Ipsenol attracts mates, as well as other males who initiate their own galleries. Females enter a nuptial chamber, mate, and excavate egg galleries branching from the nuptial chamber. I. grandicollis is polygamous, and the original nuptial chamber built by the male may become the centre of a 'Y' or 'H' shape, as two to four females (occasionally more) each excavate their own egg gallery from the common nuptial chamber. The male typically remains in his nuptial chamber throughout oviposition, where he repeatedly mates with the females and assists in removing frass from the system of phloem tunnels.

Females lay up to 50 eggs, each in an individual niche cut into the phloem tissue of the oviposition gallery wall. Niches are individually covered by the female with plugs of chewed phloem. Eggs hatch after 3-5 days, and larvae begin mining their individual feeding galleries at right angles to the egg gallery. They pass through three larval instars, each lasting several days. Pupation lasts another 3-5 days, after which the callow adults spend another 1 or 2 days within the pupal chamber; during this time the colour of the adult changes from a very pale tan to a dark or reddish brown. A period of pre-maturation feeding begins where the adult meanders through the host material phloem, feeding as it goes, for a period of up to several weeks, depending upon the geographic location and season. After this period, the sexually mature adult emerges by chewing a small, round exit hole out through the bark.

In a trial in west-central Wisconsin under natural summer temperatures (average daily mean, maximum, minimum air temperatures during the period = 19.4, 36.6, 0.2°C, respectively) the median development time for a population in a log, from attack to emergence, was 76 days (Ayres et al., 2001). The development process is highly temperature dependent, however, so this can be expected to vary widely. In the southern USA, where I. grandicollis is multivoltine, emerging adults immediately begin flights in search of a suitable host. In the north-central USA, where it is univoltine, emerging adults either go directly to an overwintering site in the forest litter layer, or to a temporary feeding site. Spring emergence from overwintering sites is apparently cued by rising soil and air temperatures in the spring. In Wisconsin, first flights typically occur when soil temperatures rise above 5°C and air temperatures exceed 15°C. The lower lethal temperature for adults is -12 to -18°C depending upon genotype and season (Lombardero et al., 2000). All immature life stages are extremely vulnerable to winter mortality, both because they tend to be less cold tolerant and because they remain within the phloem of host trees where they seldom benefit from insulation by snow (Lawson, 1993; Lombardero et al., 2000).

Within its native forests of North America, I. grandicollis forms part of a large community that co-occurs and interacts within the phloem layer of dead and dying trees (Riley and Goyer, 1988). These diverse assemblages usually include other bark beetles, several groups of insect predators and parasitoids (see Notes on Natural Enemies), numerous species of mites and fungi (commonly phoretic on adult bark beetles), wood boring insects (especially Cerambycidae and Buprestidae) and many other species that are fungivores, scavengers, or generalist predators. Major outbreaks of I. grandicollis seem to be less common within its native forests than in Australia, perhaps because population fluctuations within the rich native community are dampened by species interactions. However, pestilence in native forests may sometimes be exacerbated when I. grandicollis is able to co-operate with congeners in mass attacks of host trees (Ayres et al., 2001) or benefit from outbreaks by more aggressive bark beetles such as Dendroctonus frontalis.

Natural enemies

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Notes on Natural Enemies

Top of page In the USA, I. grandicollis has a plethora of natural enemies, primarily arthropod predators and parasitoids. The adults of several species of Cleridae prey on adult I. grandicollis during their initial colonization of host trees, and clerid larvae prey on I. grandicollis larvae during subsequent brood development within host trees. Of these, Thanasimus dubius is the most abundant and widespread. In the southern USA, T. dubius tends to be supplanted as a predator of I. grandicollis by Temnochila virescens [Temnoscheila virescens] (Trogossitidae). Several species of Histeridae, e.g. Platysoma cylindricum, are also abundant and, presumably important, predators of I. grandicollis. Dipterans, of which Medetera bistriata (Dolichopodidae) may be most significant, are voracious predators in their larval stage, prowling the bark beetle galleries feeding on larval and pupal stages. Parasitic wasps such as Roptrocerus xylophagorum (Pteromalidae) parasitize bark beetle larvae by ovipositing through the outer bark into larval feeding galleries.

Although predators and parasites have demonstrated demographic importance in controlled experiments (Riley and Goyer, 1986) and are presumably significant forces in natural populations, only one species has been found to be an effective biological control agent. Whilst Thanasimus dubius, Temnoscheila virescens and R. xylophagorum have been introduced into Australia, only R. xylophagorum has established and impacted upon I. grandicollis. R. xylophagorum is now widespread in Australia and can achieve up to 70% parasitism (Waterhouse and Sands, 2001).

Means of Movement and Dispersal

Top of page Some bark beetles are strong fliers and can migrate long distances. The most common means of introduction to new areas is by unseasoned sawn wood and wooden crates with bark on them. Debarking wood prevents the introduction of bark beetles. Most scolytids intercepted in the USA are found on dunnage, which is a difficult material to monitor (EPPO/CABI, 1997).

Plant Trade

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Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Growing medium accompanying plants
Leaves
Roots
Seedlings/Micropropagated plants
True seeds (inc. grain)
Wood

Wood Packaging

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Impact

Top of page In 2000, losses due to Ips grandicollis and two other sympatric Ips species (Ips avulsus and Ips calligraphus) were estimated to be $13.4 million annually in southern USA (SFIWC, 2001). This is probably an underestimate because many attacked trees are scattered across the landscape and overlooked in forest inventories (Riley and Goyer, 1988). Additional losses are incurred by the degradation of logs, which occurs both before and after felling, due to the blue stain defect in lumber caused by fungi of the genus Ophiostoma, for which I. grandicollis is a vector. In Australia, losses in 1994 were estimated to be several million Australian dollars following bushfires that damaged trees in 8688 ha of plantations (Wylie et al., 1999). This loss was primarily due to the blue stain degradation of trees killed by the fire, but also involved attacks of apparently healthy trees by the I. grandicollis populations that developed within the fire area (Hood and Ramsden, 1997).

Environmental Impact

Top of page Some bark beetle species can cause dramatic disturbance to forests by killing large tracts of mature pines. However, this is seldom the case with I. grandicollis. In North America, I. grandicollis should probably be regarded as a natural element in healthy forest ecosystems. I. grandicollis typically subsists in trees that have been killed or compromised by windstorms, lightning, fire, or severe drought. In this sense, the beetles are scavengers rather than agents of disturbance. Furthermore, they are arguably keystone species for hundreds of other species of arthropods, nematodes, fungi and bacteria (Savely, 1939; Ayres et al., 2001). This community contributes to the food base of some vertebrates such as salamanders (Caudata), woodpeckers (Picidae), armadillos (Dasypodidae), raccoons (Procyon spp.) and black bears (Ursus americanus). As initial colonizers of dying trees, they influence the initial ecological trajectory for decomposition of coarse, woody debris and the recycling of minerals within forest ecosystems. When attacking scattered trees, they probably promote diversity by creating openings within the forest canopy. The 'snags' that result from recently dead Pinus spp. trees frequently provide important structural habitat for assorted species of birds, bats (Chiroptera), and squirrels (Sciuridae). It is possible that interspecific competition from I. grandicollis provides some natural control against outbreaks by more aggressive species of bark beetles.

Detection and Inspection

Top of page Local presence of I. grandicollis can be monitored using funnel traps baited with the male aggregation pheromone ipsenol. However, since this species maintains an endemic presence in most pine forests within its range, its presence does not presage imminent outbreak.

Similarities to Other Species/Conditions

Top of page I. grandicollis is similar in size, colour and shape to many species of bark beetles. In adults, the deeply scooped out declivity bordered by five pairs of spines, the hidden head and the flattened, imperfectly round antennal club distinguish it from all bark beetles except for a few other species of five-spined Ips, from which it cannot easily be distinguished except by host, geographical range, or genitalia. The other life stages are virtually impossible to differentiate from many other phloem-feeding bark beetles.

Damage to trees by I. grandicollis is very similar to damage caused by other bark beetles, especially other members of the tribe Ipini. I. grandicollis often coexists with other bark beetles within the same phloem tissues of the same tree.

Prevention and Control

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Introduction

In the forests of the USA, I. grandicollis is considered the least aggressive of the bark beetle species that it co-occurs with; therefore, control measures are rarely focused on it. Within that community, it cannot be ascertained whether I. grandicollis would cause more damage in the absence of other bark beetles, or less, and in fact such a question may be irrelevant in the context of pest control. In other regions of the world, especially where it is an exotic, control measures may be more necessary, more effective, and more justifiable.

As with most pests, control of I. grandicollis is comprised of three strategies: monitoring, prevention and suppression. Control goals may be to either prevent tree mortality or to reduce blue stain damage (I. grandicollis is a vector for blue stain fungi), or both; and although similar strategies may be employed toward both goals, they may differ.

Field Monitoring

Several pheromone trap designs have been tested (Rose et al., 1981; McCravy et al., 2000) and shown to be effective for monitoring population size when baited with the aggregation pheromone ipsenol. Aerial forest surveys can readily detect and help quantify outbreaks and attacks. Such monitoring can help determine the need for suppression tactics. It is probable that the response threshold will differ depending on the situation and size and nature of the damage, but in many cases it may be the same as a natural response threshold that theoretically corresponds to the escape threshold in multiple equilibria models of population dynamics (Berryman, 1987). In the case of blue stain prevention, it may be easier and more effective to focus efforts on reducing the process time of infected material rather than to try to prevent infection.

Cultural Control and Sanitary Methods

There are several prevention tactics that may reduce the likelihood of insect populations reaching a crucial threshold. Harvesting practices like slash sanitation during logging, prompt movement of logs from logging sites, and prompt salvage logging after natural disasters like fires and windstorms may help prevent population growth to dangerous levels. Silvicultural practices that reduce or avoid large acreages of overstocked mono-specific stands of susceptible hosts can help, as can management plans that help maintain healthy populations of natural enemies.

Some of the same tactics for prevention can be used for suppression. Removal of infected logs in infested forest areas, salvage logging and debarking or processing of infested logs can reduce populations and minimize both unintended tree mortality and blue stain damage to logs both in the forest and in the mill yard, especially if done promptly.

Mass trapping with the same traps and lures used for monitor trapping can be effective at capturing large numbers of insects; and multiple equilibria models (Berryman, 1987) suggest that it may be possible to suppress populations below the escape threshold. Success has not been demonstrated, however, and whether such mass trapping campaigns decrease populations enough to reduce damage remains arguable.

Chemical Control

Many synthesized and naturally occurring chemical substances have been tested for use as possible prevention and suppression tactics. One strategy involves using semiochemical disruption of the aggregation process. Certain green leaf volatiles (Dickens et al., 1992), 4-allylanisole (Hayes et al., 1996), verbenone, pine oil (Berisford et al., 1986; Nord et al., 1990) and a wide range of odoriferous compounds (All and Anderson, 1974) have been tested. Of these, a few showed some potential (verbenone, green leaf volatiles, ammonium hydroxide and 4-allylanisole), but none have become cost-effective solutions.

Many other chemicals have been tested as insecticides, such as carbaryl, chlorpyrifos-methyl, deltamethrin and propoxur (Ragenovich and Coster, 1974; Stone and Simpson, 1987). Fuel oil mixtures have been shown to cause high I. grandicollis mortality when applied on logs (Cibulsky and Hyche, 1974). Of these, only carbaryl, under the trade name Sevimol, is currently labelled for use in the USA, along with permethrin (trade name Astro) for control of bark beetles. There is also a pending registration for bark beetle control for bifenthrin (trade name Onyx).

It must be noted that both insecticides and deterrents have limited utility due to the cost of application and degree and duration of protection, and have never been shown to be effective in the forest setting. Additionally, they may have side effects that may outweigh possible benefits. They may, however, have limited use in protecting individual high value trees, or for temporary protection against log deck degradation.

Biological Control

Many potential candidates have been evaluated for use as biological control agents in Australia (Berisford, 1989). Neither of the introduced predators, Temnoscheila virescens and Thanasimus dubius, are known to be established, but two parasitoids have established. Of these, only Rhoptocerus xylophagorum has had any impact. R. xylophagorum is now widespread and causes up to 70% parasitism. This attack, combined with much improved silvicultural management (including removal of bark from slash and logs), has greatly reduced the damage caused by I. grandicollis (Waterhouse and Sands, 2001).

References

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Abbott I, 1993. Review of the ecology and control of the introduced bark beetle Ips grandicollis (Eichoff) (Coleoptera): Scolytidae in Western Australia, 1952-1990. CALMScience, 1(1):35-46; 29 ref.

All JN, Anderson RF, 1974. The influence of various odors on host selection by pioneer beetles of Ips grandicollis. Journal of the Georgia Entomological Society, 9(4):223-228

Anderson RF, 1977. Dispersal and attack behavior of the southern pine engraver, Ips grandicollis Eichh., Coleoptera, Scolytidae. In: Kulman HM, Chiang HC, ed. Technical Bulletin, University of Minnesota Agricultural Experiment Station, 17-23

Arnold DC, Jiracek SR, 1982. Beetles of Oklahoma Pines. Publication No. MP-113. Agricultural Experiment Station, Oklahoma State University Stillwater, Oklahoma USA, 16 pp.

Ayres BD, Ayres MP, Abrahamson MD, Teale SA, 2001. Resource partitioning and overlap in three sympatric species of Ips bark beetles (Coleoptera : Scolytidae). Oecologia, 128(3):443-453.

Berisford CW, 1989. Biological control of pine bark beetles: new approaches to an old problem. In: Entomology in Virginia: New Problems and New Approaches, held in Virginia, USA, on 8 September 1989.

Berisford CW, Brady UE, Fatzinger CW, Ebel BH, 1986. Evaluation of a repellent for prevention of attacks by three species of southern pine bark beetles (Coleoptera: Scolytidae). Journal of Entomological Science, 21(4):316-318

Berrios C, Menendez JM, Rodriguez M, 1987. Presence of Ips on new species of pines in the north of Matanzas province. Revista Forestal Baracoa, 17(2):113-115

Berryman AA, 1987. The theory and classification of outbreaks. In: Barbosa P, Schultz JC, eds. Insect Outbreaks. San Diego, USA: Academic Press, Inc., 3-30.

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, 2007. Ips grandicollis. [Distribution map]. Distribution Maps of Plant Pests, No.June. Wallingford, UK: CABI, Map 691.

Coster JE, 1974. Evaluation of some carbamate and phosphate insecticides against southern pine beetle and Ips bark beetles. Journal of Economic Entomology, 67(6):763-765.

Dickens JC, Billings RF, Payne TL, 1992. Green leaf volatiles interupt aggregation pheromone response in bark beetles infesting southern pines. Experientia, 48(5):523-524; 21 ref.

Garraway E, 1986. The biology of Ips calligraphus and Ips grandicollis (Coleoptera: Scolytidae) in Jamaica. Canadian Entomologist, 118(2):113-121

Garraway E, Freeman BE, 1990. The population dynamics of Ips grandicollis (Eichhoff) (Coleoptera: Scolytidae) in Jamaica. Canadian Entomologist, 122(3-4):217-227

Guerra C, Lopez MO, Valdes E, Perez P, Fernandez A, 1989. Presence of Ceratocystiopsis minima and Ceratocystis seticollis in pine stands in the western region of Cuba. Revista Forestal Baracoa, 19(2):113-117

Haack RA, Paiz-Schwartz G, 1997. Bark beetle (Coleoptera: Scolytidae) outbreak in pine forests of the Sierra de las Minas Biosphere Reserve, Guatemala. Entomological News, 108(1):67-76; 40 ref.

Hayes JL, Meeker JR, Foltz JL, Strom BL, 1996. Suppression of bark beetles and protection of pines in the urban environment: a case study. Journal of Arboriculture, 22(2):67-74; 16 ref.

Hayes JL, Strom BL, Roton LM, Ingram LL Jr, 1994. Repellent properties of the host compound 4-allylanisole to the southern pine beetle. Journal of Chemical Ecology, 20(7):1595-1615

Hoffard WH, Coster JE, 1976. Endoparasitic nematodes of Ips bark beetles in eastern Texas. Environmental Entomology, 5(1):128-132

Hood IA, Ramsden M, 1997. Sapstain and decay following fire in stands of Pinus elliottii var. elliottii near Beerburrum, south east Queensland. Australian Forestry, 60(1):7-15; 14 ref.

Kinn DN, 1984. Incidence of endoparasitic nematodes in Ips engraver beetles in central Louisiana. Journal of the Georgia Entomological Society, 19(3):322-326

Lawson SA, 1993. Overwintering mortality of Ips grandicollis Eichh. (Col., Scolytidae) and its parasitoid, Roptrocerus xylophagorum Ratz. (Hym., Pteromalidae), in South Australia. Journal of Applied Entomology, 115(3):240-245

Lombardero MJ, Ayres MP, Ayres BD, Reeve JD, 2000. Cold tolerance of four species of bark beetle (Coleoptera: Scolytidae) in North America. Environmental Entomology, 29(3):421-432; 88 ref.

McCravy KW, Nowak JT, Douce GK, Berisford CW, 2000. Evaluation of multiple-funnel and slot traps for collection of southern pine bark beetles and predators. Journal of Entomological Science, 35(1):77-82; 20 ref.

Morgan FD, 1967. Ips grandicollis in South Australia. Australian Forestry, 31:137-155.

Morgan FD, 1989. Forty years of Salix noctilio and Ips grandicollis in Australia. New Zealand Journal of Forestry Science, 19(2-3):198-209

Neumann FG, Marks GC, 1990. Status and management of insect pests and diseases in Victorian softwood plantations. Australian Forestry, 53(2):131-144; 49 ref.

Neumann FG, Morey J, 1984. Studies on the introduced bark beetle Ips grandicollis (Eichhoff) in Victorian radiata pine plantations. Australian Forest Research, 14(4):283-300

Nord JC, Hastings FL, Jones AS, 1990. Field tests of pine oil as a repellent for southern pine bark beetles. Research Note - Southeastern Forest Experiment Station, USDA Forest Service, No. SE-355:8 pp.

Nunberg M, 1974. The occurrence of Ips grandicollis (Eichh.) (Coleoptera, Scolytidae) in Cuba. Polskie Pismo Entomologiczne, 44(4):735-736

Ragenovich IR, Coster JE, 1974. Evaluation of some carbamate and phosphate insecticides against southern pine beetle and Ips bark beetles. Journal of Economic Entomology, 67(6):763-765

Renwick JAA, Vite JP, 1972. Pheromones and host volatiles that govern aggregation of the six-spined engraver beetle, Ips calligraphus. Journal of Insect Physiology, 18(6):1215-1219.

Riley MA, Goyer RA, 1986. Impact of beneficial insects on Ips spp. (Coleoptera: Scolytidae) bark beetles in felled loblolly and slash pines in Louisiana. Environmental Entomology, 15(6):1220-1224

Riley MA, Goyer RA, 1988. Seasonal abundance of beneficial insects and Ips spp. engraver beetles (Coleoptera: Scolytidae) in felled loblolly and slash pines in Louisiana. Journal of Entomological Science, 23(4):357-365

Rimes GD, 1959. The bark beetle in West Australian Pine forests. Journal of Agriculture of Western Australia, Perth, (Ser. 3) 8(3).

Rose WF III, Billings RF, Vite JP, 1981. Southern pine bark beetles: evaluation of non-sticky pheromone trap designs for survey and research. Southwestern Entomologist, 6(1):1-9

Savely HE, 1939. Ecological relations of certain animals in dead pine and oak logs. Ecological Monographs, 9:321-385.

Schmitt JJ, Goyer RA, 1983. Consumption rates and predatory habits of Scoloposcelis mississippensis and Lyctocoris elongatus (Hemiptera: Anthocoridae) on pine bark beetles. Environmental Entomology, 12(2):363-367

SFIWC, 2001. Report on losses caused by forest insects. In: 45th Southern Forest Insect Work Conference, Georgia, USA, 23-26 July.

Smith IM, McNamara DG, Scott PR, Holderness M, 1997. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization. Quarantine pests for Europe. Second Edition. Data sheets on quarantine pests for the European Union and for the European and Mediterranean Plant Protection Organization., Ed. 2:vii + 1425 pp.; many ref.

Stone C, 1990. Parasitic and phoretic nematodes associated with Ips grandicollis (Coleoptera: Scolytidae) in New South Wales, Australia. Nematologica, 36(4):478-480

Stone C, Simpson JA, 1987. Influence of Ips grandicollis on the incidence and spread of bluestain fungi in Pinus elliottii billets in north-eastern New South Wales. Australian Forestry, 50(2):86-94

Stone C, Simpson JA, 1990. Species associations in Ips grandicollis galleries in Pinus taeda. New Zealand Journal of Forestry Science, 20(1):75-96; 30 ref.

Waterhouse DF, Sands DPA, 2001. Classical biological control of arthropods in Australia. Canberra, Australia: Australian Centre for International Agricultural Research (ACIAR), 559 pp.

Werner RA, 1972. Aggregation behavior of the beetle Ips grandicollis in response to (1) host-produced attractants; (2) insect-produced attractants. Journal of Insect Physiology, 18:423-437.

Wood SL, 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs, No. 6:1359 pp.

Wylie FR, Peters B, DeBaar M, King J, Fitzgerald C, 1999. Managing attack by bark and ambrosia beetles (Coleoptera:Scolytidae) in fire-damaged Pinus plantations and salvaged logs in Queensland, Australia. Australian Forestry, 62:148-153.

Distribution Maps

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