Preferred name: Monochamus carolinensis
Authority: Olivier
Taxonomic position: Animalia: Arthropoda: Hexapoda: Insecta: Coleoptera: Cerambycidae
Common names in English: carolina sawyer
view more common names online...
EPPO Categorization: A1 list
view more categorizations online...
EPPO Code: MONCCA

Monochamus carolinensis only attacks and colonizes the members of genus Pinus, including jack pine (P. banksiana), red pine (P. resinosa), eastern white pine (P. strobus) (Linsley & Chemsak, 1997), Scots pine (P. sylvestris) (MacRae, 1993, Pershing & Linit, 1986b) shortleaf pine (P. echinata) (Walsh & Linit, 1984; MacRae, 1993), loblolly pine (Pinus taeda) (Alya & Hain, 1985). During experiments, adults of M. carolinensis were trapped by flight traps placed at the mid-bole of stressed Virginia pines (Pinus virginiana) (Hines & Heikkenen, 1977) showing that they were attracted by these trees, but no adult beetles emerged from the trees.

Host list: Pinus banksiana, Pinus echinata, Pinus resinosa, Pinus strobus, Pinus sylvestris, Pinus taeda

Monochamus carolinensis is native to North America and occurs from South-Eastern Canada (Ontario, Quebec, and New Brunswick) through the eastern half of the USA, from Texas to Minnesota. There is only one record from Mexico (Durango) by Dillon & Dillon (1941).

North America: Canada (New Brunswick, Ontario, Québec), Mexico, United States of America (Alabama, Arkansas, Connecticut, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Nebraska, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Virginia, West Virginia, Wisconsin)

The development of species in the genus Monochamus takes several months to two years (Webb, 1909, Rose, 1957; Akbulut et al., 2017). The life cycle of M. carolinensis appears to be temperature dependent. In the USA, two-year generations are common in the northern range and two generations per year in the southern range of M. carolinensis (Pershing & Linit, 1986a). Adult beetles emerge through circular holes cut in the bark and fly to healthy pine trees, feed on the bark and phloem of twigs and branches prior to mating and oviposition (maturation feeding) for 10-14 days (Akbulut & Stamps, 2012). It is at this point that pine wood nematodes (Bursaphelenchus xylophilus), if present on the beetle, exit the beetle’s trachea and enter through feeding wounds in the bark. During the night, beetles fly to dead or stressed pines to mate and oviposit in response to volatile chemicals released by trees. After mating, the female deposits an egg below the bark (Walsh & Linit, 1985). If the pine wood nematodes are present in the bodies of female beetles, they may enter oviposition wounds at this time and feed on fungus present in the stressed or dying pine. The number of eggs deposited in an oviposition wound differs both within and among Monochamus species (Akbulut et al., 2017). The oviposition wounds can be slit like on thin bark or pit like on thick bark, and contain one egg in most cases (Walsh & Linit, 1985), eggless, or multiple eggs can be laid in a single wound. The egg incubation period ranges from 6 to 9 days under natural conditions but can vary from 10 to 12 days according to temperature (Kobayashi et al., 1984). The egg hatches and the early instars develop entirely in the subcortical zone of the host trees (Pershing & Linit, 1986b). Larvae can feed on inner bark, cambium, and outer sapwood (Linit, 1988). Late instar larvae construct galleries in the sapwood. Each larva constructs its own gallery. M. carolinensis has three to eight larval instars prior to pupation (Pershing & Linit, 1986a). The insect pupates in a chamber in the woody tissue near the cambium layer (Linit, 1988). The adult ecloses within the pupal chamber and remains there for several days until cuticular tanning is complete before exiting the tree. At this point, if present, the dispersal form of the pinewood nematode exits the surrounding wood tissue, crawls onto the body of the insect, and enters the spiracles and trachea for transportation to new host trees. The beetle overwinters most commonly as a larva, but eggs and pupae overwinter as well (Pershing & Linit, 1986a). Adult emergence, which occurs by chewing a round exit hole in the wood and bark, starts in the spring and continues until the late summer. The total development time from oviposition to emergence averages 8-12 weeks (Pershing & Linit, 1986b). In the laboratory, when reared with Pinus sylvestris logs, the within-wood developmental period of M. carolinensis ranges from 83 to 108 days under constant conditions (Akbulut & Linit 1999a, 1999b). Linsley (1961) reported that duration of the larval stages of cerambycids can be shorter when they feed on a freshly killed host and longer when they feed on wood that has been dead for some time.

Several studies have examined the life history traits of M. carolinensis. Previous laboratory studies (Linit, 1985; Akbulut et al., 2004) reported that within-log generation survival of M. carolinensis ranged from 6 % to 15 %. Alya & Hain (1985) found within-log generation survival of M. carolinensis and M. titillator to be 15 % in logs maintained in outdoor cages in North Carolina. The high mortality of beetles within the logs suggested that either intraspecific competition among M. carolinensis larvae and/or interspecific competition among M. carolinensis and other beetle species occurred within the logs. Akbulut et al. (2004) examined the relationships between initial oviposition density and log volume, bark area, and adult survival. They found that the number of adults emerged per log was not related to the number of eggs laid. Late larval and adult numbers were significantly correlated with log size, and only 12% of the initial cohort completed development and emerged as adults. Akbulut et al. (2004) attributed the high within-log mortality of beetles to intraspecific competition and cannibalism.

There is a great variation in the longevity of Monochamus beetles (Akbulut et al., 2017). M. carolinensis adults that emerge from P. sylvestris logs cut in autumn and spring live for 38 and 59 days, respectively, under laboratory conditions (Akbulut & Linit, 1999b). Togashi et al. (2009) reported that the mean longevity of males and females in M. carolinensis is 51 and 56 days respectively in the field cages. Similar variations were reported for M. galloprovincialis (Naves et al., 2006; Akbulut, 2009) and M. alternatus (Togashi & Magira 1981; Togashi, 1997). Variations in adult longevity can be attributed to the nutritional quality of the phloem and cambium of pine logs consumed at the larval stage, and also to the quality of the twig subcortical tissues consumed at the adult stage (Akbulut & Linit, 1999b).

The total number of eggs laid by a female (fecundity) varies among species and within species. For example, it ranges from 0 to 384 for M. carolinensis (Walsh & Linit, 1985; Akbulut, 1998), and from 0 to 343 for M. alternatus (Kobayashi et al., 1984; Kobayashi, 1988; Togashi, 1997). M. galloprovincialis laid an average of 67 eggs in P. pinaster in Portugal (Naves et al., 2006), 138 in P. sylvestris in France (Koutroumpa et al., 2008), 57 eggs in P. nigra in Türkiye (Akbulut, 2009).

Monochamus spp. beetles are highly attracted to dead or stressed pine trees. Pheromones for sex attraction may not exist because adult beetles of both sexes are brought together by attraction to volatiles released by the physiological breakdown of pine trees. M. carolinensis males search for females (Edwards & Linit, 1991). No sex pheromone is known to exist, and antennal contact between sexes is sufficient for mate recognition in M. carolinensis. However, several studies have suggested the existence of sex or aggregation pheromones for other Monochamus species (Akbulut et al., 2017). Two types of pheromones have been reported for M. alternatus: 1) a male sex pheromone that attracts the female and 2) a contact pheromone on the body surface of the female that elicits mate recognition and initiation of courtship (Fauziah et al., 1987; Kim et al., 1992). Ibeas et al. (2008) reported that mature female M. galloprovincialis under laboratory conditions were attracted to volatile compounds produced by mature male beetles; however, the males did not respond to female-produced volatiles. A chemical substance of 2-undecyloxy-1-ethanol (monochamol) has been proven to be a male produced aggregation pheromone for M. galloprovincialis (Pajares et al., 2010), M. alternatus (Teale et al., 2011), M. carolinensis and M. titillator (Allison et al., 2012, Rasatti et al., 2012), M. scutellatus scutellatus (Fierke et al., 2012) and M. sutor (Pajares et al., 2013). Macias-Samano et al., (2012) suggest that monochamol is a likely pheromone component for M. clamator and M. obtusus and attractant for M. notatus (Fierke et al., 2012). Ryall et al. (2015) proved that monochamol is attractive to 12 Monochamus species including M. carolinensis and has a good potential for surveys of the genus Monochamus.

Monochamus beetles are generally poor fliers, even though in flight mill experiments, a M. carolinensis beetle has flown a maximum distance of 10 km with a duration of 115 minutes (Akbulut & Linit, 1999c). In the same study, the mean flight distance was reported to be 2.2 km for 1-day-old M. carolinensis adult beetles. In mark and recapture studies of the closely related beetle, M. alternatus, Ogawa & Hagiwara, (1980) found that most of the beetles were recaptured within 100 m of the release site, whereas Kobayashi et al. (1984) reported that beetles mostly moved 800 m from their origin and some flew up to 3.3 km. However, there are no flight performance studies under field conditions for M. carolinensis. Mas et al. (2013) carried out mark-release-recapture trials with M. galloprovincialis and marked adults flew a maximum distance of 22.1 km and about 2% of released beetles travelled more than 3.0 km. Another mark-recapture study carried on M. galloprovincialis found that the lifetime adult dispersal distance was 107-122 m on average with a maximum of 464 m (Torres-Villa et al., 2015). Several studies were carried out to analyze the spread of pine wilt disease in different countries. The yearly spread of pine wilt disease (due to vector movements) was an average of 7.5 km/year in China (Robinet et al., 2009), an average of 6.609km/year in Japan (Osada et al., 2018) and an average of 5.3 km with a maximum of 8.3 km in Portugal (de la Fuente et al., 2018). The spread capacities of different Monochamus species seem to allow a few kilometres in a single flight and cumulatively reach over several tens of kilometres during the life span of a beetle (EFSA, 2018).

Symptoms

The following symptoms and signs may be related to the presence of Monochamus spp. (Wilson, 1975, Akbulut & Stamps, 2012):

1. The presence of slit like or pit like oviposition scars chewed into the bark of pine trees.
2. The presence of feeding tracks and oval shaped larval entrance holes in the sapwood caused by larval feeding when the bark is removed.
3. Excelsior-like frass around cut logs from holes in the bark or at the base of trees colonized by the beetle.
4. A round exit hole in the wood and bark created by adults.

Morphology

Egg

Eggs are white, elongate, cylindrical, and slightly flattened with rounded ends. The average size is 2.5 - 3.0 mm long by 0.9 mm wide (CABI, 2019).

Larva

Larvae are soft-bodied, elongate, dirty white in colour, with a light-yellow thorax and amber-brown head. Larvae have ten abdominal segments, are legless, and have a pair of short, stout mandibles. The head is longer than it is wide. Mature larvae may be 25 - 50 mm in length (CABI, 2019).

Pupa

Pupae are initially white and resemble adults in body form. They are 20 -25 mm in length, with antennae tightly curled in spirals flat against the sides of the body. The legs, antennae and wings project from but are closely held to the body. As the pupa matures, it darkens and the limbs become more distinct (CABI, 2019).

Adult

Adult beetles are fairly large, dark reddish-brown insects, 13 - 25 mm in length and 5 -7 mm in width. Male antennae are two times the length of the body, and female antennae are approximately equal to or slightly longer than the length of the body. The antennae are high on the head, close together, and separated by a deep hollow. Males may be slightly smaller and more slender in body shape than females. Both sexes possess a stout pair of mandibles and long, thin legs. The elytra are dark reddish brown with three indistinct bands of white pubescence, giving the elytra a mottled, spotted dirty-white and brown appearance. Each side of the pronotum has a large lateral submedian conical protuberance. The elytra are wider than the pronotum at the base (CABI, 2019).

Detection and inspection methods

Monochamus spp. are attracted to weakened, dying, dead, or freshly cut trees. Therefore, such trees, which often have partly or completely discoloured needles, should be the focus of surveillance for Monochamus spp. Close inspection of these trees may allow the detection of oviposition scars in the bark, oval shaped larval entrance holes in the sapwood under the bark or round adult emergence holes in the sapwood. Other cerambycids make similar oviposition scars and emergence holes, so their presence is not a certain confirmation of M. carolinensis. Excelsior-like frass may be present around cut logs, from holes in the bark, or at the base of trees infested with the beetle. The cryptic nature of the beetle and its camouflage coloration make it difficult to observe on trees in a natural setting. However, adults can be seen walking or resting on cut and dead wood (EFSA, 2018).

Monochamus beetles are considered to be poor fliers, although there is evidence in the literature (details given in the biology section) that they can perform both short-distance and long-distance dispersals (Akbulut et al., 2017). According to flight mill, field experiments and models developed Monochamus species can naturally disperse by flight a few kilometres to tens of kilometres during their life span. The human assisted spread of Monochamus spp. may cover hundreds or thousands of kilometres. Larvae of Monochamus spp. develop entirely within the wood, and they are difficult to detect. Therefore, they can be easily transported as untreated conifer wood products in trade. Monochamus spp. can be spread by coniferous wood (round or sawn) and wood packaging materials, dunnage from conifers, particle and waste wood of host trees, coniferous wood products and as hitchhikers in containers (EFSA, 2018, 2020). Plants for planting are not likely to be the pathway for the spread of M. carolinensis, as they attack dead or stressed pine trees.

Economic importance

Monochamus spp. beetles are highly attracted to dead, stressed or recently cut coniferous trees. Therefore, many of them are considered secondary pests, and typically they do not cause the death of trees. M. carolinensis may do little damage to healthy trees in the absence of the pine wood nematode (Akbulut & Stamps, 2012). Pine wilt disease is rarely the cause of mortality in native conifers in North America because of the coevolution of the native Monochamus species and tree species (Akbulut & Stamps, 2012). The pine wood nematode is widely distributed in the USA, and the disease is most serious in pine trees that are exotic to North America, such as Scots, Austrian, Japanese red and Japanese black pines. Beetles transmit nematodes from dead trees to healthy trees through feeding wounds of newly emerged beetles (during maturation feeding) and weakened, dying, or recently cut trees through oviposition wounds (during egg oviposition). Infection of trees with nematodes under favourable conditions leads to the mass mortality of susceptible pine stands. The disease has a devastating impact on native pine forests in Japan, China, Korea, Taiwan, and Portugal. In the USA, the disease affects exotic pines planted on private properties and natural areas planted with susceptible, non-native, and poorly adapted pine species. Scots pine used in windbreaks, landscape plantings, and Christmas tree farms and nurseries can be affected by pine wilt disease. For lumber producers, timber quality and value can be lessened by beetle’s presence due to larval galleries and the blue stain fungi that are often associated with the beetle's presence in wood (CABI, 2019). Pine wilt disease has a substantial impact on the international wood trade from countries where the nematode and its vector are present to the European Union and other countries. Economic loss as a result of trade restrictions on softwood products is difficult to measure due to environmental, regulatory and supply issues (CABI, 2019), but has been discussed in several publications (Dwinell, 1997; Anon., 1991; Bergdahl, 1988, Miller et al., 2013). 

Control

In North America, M. carolinensis and the pine wood nematode are native and widely distributed. Therefore, control of M. carolinensis is usually limited to the removal and burning of infested trees and the timely spraying of insecticides on susceptible exotic pines such as Pinus sylvestris (i.e., on Christmas tree plantations). Prevention is the best method for controlling beetles. Adult beetles are attracted to recently dead, dying, or freshly cut trees, so dead and dying trees must be cut down and bark removed from felled trees as quickly as possible to eliminate breeding habitat for the beetles and to prevent nematode transmission to healthy trees (Donald et al., 2016). The stumps of logged trees should also be removed or buried as they are also attractive to M. carolinensis. Monochamus damage can also be prevented by not exposing wood during the egg-laying period of beetles (Dwinell, 1997; Gibson, 2010). Persons in North America who have infected exotic pines on their properties have to remove the infected trees, burn them (CABI, 2019), and are encouraged to plant resistant native conifer species instead of susceptible tree species as ornamentals. Existing valuable trees can be protected from pine wilt disease with preventive injections of nematicide (Donald et al., 2016).

Pheromones and traps play an important role in monitoring and controlling M. carolinensis. A recent study proved that monochamol (male aggregation pheromone) has good potential for surveying the genus Monochamus including M. carolinensis (Ryall et al., 2015). The blend of monochamol, ipsenol methyl-butenol and α-pinene used with black Teflon-coated cross-vane traps proved to be an efficient attractant to several Monochamus species (Boone et al., 2018).

The use of biological control agents against M. carolinensis has not been studied (CABI, 2019). The fungus Beauveria bassiana has been found to be effective against the closely related M. alternatus (Shimazu et al., 1995; Shimazu, 2009).

Phytosanitary risk

Accidental introduction of M. carolinensis together with the pine wood nematode to new locations increases the rate of spread of the pine wood nematode. The risk of introduction of both pests into the EPPO region is high (Evans et al., 2009; EPPO, 2022). M. carolinensis is known to be the main vector of B. xylophilus in North America (Akbulut & Stamps, 2012). The egg, larval, and pupal stages of the beetle can be transported via unprocessed softwood logs. Introduction of pine wood nematode-infected Monochamus species can be limited by import requirements specified for host plants of the pine wood nematode and coniferous woods by National or Regional Plant Protection Organizations. Monochamus species are regularly intercepted in the European Union (EFSA, 2018). For example, according to the EUROPHYT database between 1999 and 2019, there have been 21 interception records of M. alternatus, non-European Monochamus species, on wood packaging material or dunnage (EFSA, 2020).

Accidental introduction of M. carolinensis and the pine wood nematode to the EPPO region can be limited by specific import requirements for host plants of the pine wood nematode and its vectors. Due to the cryptic life stages of M. carolinensis, visual inspection of timber does not always reveal the presence of eggs, larvae, or pupae under the bark or in the wood of imported products. Therefore, international standards have been developed to prevent the importation of infested wood products (FAO, 2017, EPPO 2018a, b). The phytosanitary measures recommended by the EPPO commodity standard for Coniferae, PM 8/2 (3) (EPPO, 2018b) are considered to be effective to prevent the introduction and spread of non-European Monochamus spp. and the pine wood nematode. There are recommendations for wood from Pinus, originating in countries where M. carolinensis is present: it should originate from a pest free area and be transported outside of M. carolinensis flight period, or the wood should be covered during the transportation (EPPO, 2018b). Following quarantine treatments for conifer wood are recommended: wood should be bark-free, and either heat treated according to EPPO Standard PM 10/6 or treated with ionizing radiation according to EPPO Standard PM 10/8 or appropriate fumigation and details specified on the Phytosanitary Certificate (EPPO, 2018b). The treatment of wood packaging material, including dunnage, should meet the requirements of ISPM no. 15 (FAO, 2019) to prevent the introduction of B. xylophilus and Monochamus spp. via international trade into uninfested areas.

Akbulut S (1998) Effect of Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae) Fourth stage dispersal juveniles and log seasonality on life processes of Monochamus carolinensis (Coleoptera: Cerambycidae). Ph.D. Dissertation, Department of Entomology, University of Missouri-Columbia, USA.

Akbulut S (2009) Comparison of the reproductive potential of Monochamus galloprovincialis Olivier (Coleoptera: Cerambycidae) on two pine species under laboratory conditions. Phytoparasitica 37, 125-135.

Akbulut S & Linit MJ (1999a) Reproductive potential of Monochamus carolinensis (Coleoptera: Cerambycidae) with respect to nematode phoresis. Environmental Entomology 28, 407-411.

Akbulut S & Linit MJ (1999b) Seasonal effect on reproductive performance of Monochamus carolinensis (Coleoptera: Cerambycidae) reared in pine logs. Journal of Economic Entomology 92, 631-637.

Akbulut S & Linit MJ (1999c) Flight performance of Monochamus carolinensis (Coleoptera: Cerambycidae) with respect to nematode phoresis and beetle characteristics. Environmental Entomology 28, 1014-1020.

Akbulut S & Stamps WT (2012) Insect vectors of the pinewood nematode: a review of the biology and ecology of Monochamus species. Forest Pathology 42, 89-99.

Akbulut S, Stamps WT & Linit MJ (2004) Population dynamics of Monochamus carolinensis (Coleoptera: Cerambycidae) under laboratory conditions. Journal of Applied Entomology 128, 17-21.

Akbulut S, Togashi K & Linit MJ (2017) Cerambycids as Plant Disease Vectors with Special Reference to Pine Wilt. In: Wang Q (ed.). Cerambycidae of the World: Biology and Pest Management. Taylor & Francis, Abingdon, UK, pp. 209-252.

Allison JD, McKenney JL, Millar JG, McElfresh JS, Mitchell RF & Hanks LM (2012) Response of the woodborers Monochamus carolinensis and Monochamus titillator (Coleoptera: Cerambycidae) to known cerambycid pheromones in the presence and absence of the host plant volatile alpha-pinene. Environmental Entomology 41, 1587-1596.

Alya AB & Hain FP (1985) Life histories of Monochamus carolinensis and M. titillator (Coleoptera: Cerambycidae) in the Piedmont of North Carolina. Journal of Entomological Science 20, 390-397.

Anon (1991) Pinewood nematode control targeted as green lumber exports to EC given reprieve. Victoria, BC, Canada: Information Forestry, Forestry Canada, Pacific Forestry Centre, (Fall/winter), 6.

Bergdahl DR (1988) Impact of pinewood nematode in North America: present and future. Journal of Nematology 20, 260-265.

Boone C, Sweeney J, Silk P, Hughes C, Webster RP, Stephen F, Maclauchlan L, Bentz B, Drumont A, Zhao B, Berkvens N, Casteels H & Gregoire J-C (2018) Monochamus species from different continents can be effectively detected with the same trapping protocol. Journal of Pest Science 92, 3–11. https://doi.org/10.1007/s10340-018-0954-4

CABI (2019) Monochamus carolinensis CABI Compendium https://doi.org/10.1079/cabicompendium.34720

de la Fuente B, Saura S, Beck PSA & Fortin M-J (2018) Predicting the spread of an invasive tree pest: The pine wood nematode in Southern Europe. Journal of Applied Ecology 5, 2374-2385.

Dillon LS & Dillon ES (1941) The tribe Monochamini in the western hemisphere (Coleoptera: Cerambycidae). Reading Public Museum and Art Gallery Scientific Publications Number 1.

Donald PA; Stamps WT, Linit MJ & Todd TC (2016) Pine Wilt Disease. The Plant Health Instructor; The American Phytopathological Society, New York, NY, USA.

Dwinell LD (1997) The pinewood nematode: regulation and mitigation. Annual Review of Phytopathology 35, 153-166.

Edwards O R & Linit MJ (1991) Oviposition behavior of Monochamus carolinensis (Coleoptera: Cerambycidae) infested with the pinewood nematode. Annals of Entomological Society of America 84, 319-323.

EFSA (2018) Pest categorisation of non-EU Monochamus spp. EFSA Journal 16, 5435.

EFSA (2020) Pest survey card on non-European Monochamus spp. EFSA Supporting publications 2020-EN-1781. https://doi.org/10.2903/sp.efsa.2020.EN-1781

EPPO (2018a) EPPO Standards. National regulatory control systems. PM 9/1 (6) Bursaphelenchus xylophilus and its vectors: procedures for official control. EPPO Bulletin 48, 503-515.

EPPO (2018b) EPPO Standards. Commodity-specific phytosanitary measures. PM 8/2 Coniferae. EPPO Bulletin 48, 463-494.

EPPO (2022) Bursaphelenchus xylophilus. EPPO datasheets on pests recommended for regulation. https://gd.eppo.int/taxon/BURSXY/datasheet

Evans H, Kulinich O, Magnusson C, Robinet C & Schroeder T (2009) Report of a Pest Risk Analysis for Bursaphelenchus xylophilus 09/15450, 1-17. https://gd.eppo.int/taxon/BURSXY/documents

FAO (2017) ISPM 39 International movement of wood. IPPC Secretariat, FAO, Rome, 20 pp.

FAO (2019) ISPM 15. Regulation of wood packaging material in international trade. FAO, Rome, 24 pp. https://www.ippc.int/en/publications/640/

Fauziah BA, Hidaka T & Tabata K (1987) The reproductive behavior of Monochamus alternatus Hope (Coleoptera:Cerambycidae). Applied Entomology and Zoology 22, 272-285.

Fierke MK, Skabeikis DD, Millar JG, Teale SA, McElffresh JS & Hanks LM (2012) Identification of a male-produced aggregation pheromone for Monochamus scutellatus scutellatus and an attractant for the congener Monochamus notatus (Coleoptera: Cerambycidae). Journal of Economic Entomology 105, 2029-2034.

Gibson K (2010) Management guide for sawyer beetles. Forest Health Protection and State Forewsy Organisations. Available online: https://www.fs.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5187547.pdf

Hines JW & Heikkenen HJ (1977) Beetles attracted to severed Virginia pine (Pinus virginiana Mill.). Environmental Entomology 6, 123-127.

Ibeas F, Diez JJ & Pajares JA (2008). Olfactory sex attraction and mating behaviour in the pine sawyer Monochamus galloprovincialis (Coleoptera: Cerambycidae). Journal of Insect Behavior 21, 101-110.

Kim G, Takabayashi J & Tabata K (1992) Function of pheromones in mating behavior of the Japanese pine sawyer beetle, Monochamus alternatus Hope. Applied Entomology and Zoology 27, 489–497.

Kobayashi F (1988) The Japanese pine sawyer. In Dynamics of Forest Insect Populations: Patterns, Causes, Implications, ed. A. A. Beryman, 431-454. Plenum Pres. New York and London.

Kobayashi F, Yamane A & Ikeda T (1984) The Japanese pine sawyer beetle as the vector of pine wilt disease. Annual Review of Entomology 29, 115-135.

Koutroumpa FA, Vincent B, Roux-Morabito G, Martin C & Lieutier F (2008) Fecundity and larval development of Monochamus galloprovincialis (Coleoptera: Cerambycidae) in experimental breeding. Annals of Forest Science 65, 707 (pp.11).

Linit MJ (1985) Continuous laboratory culture of Monochamus carolinensis with notes on larval development. Annals of Entomological Society of America 78, 212-213.

Linit MJ (1988) Nematode-vector relationships in the pine wilt disease system. Journal of Nematology 20, 227-235.

Linsley EG & Chemsak JA (1997) The Cerambycidae of North America, part VIII: bibliography, index, and host plant index. University of California Publications in Entomology 117, 534 pp.

Macias-Samano JE, Wakarchuk D, Millar JG & Hanks LM (2012) 2-Undecyloxy-1-ethanol in combination with other semiochemicals attracts three Monochamus species (Coleoptera: Cerambycidae) in British Columbia. The Canadian Entomologist 144, 764-768.

MacRae TC (1993) Annotated checklist of the longhorned beetles (Coleoptera: Cerambycidae and Disteniidae) occurring in Missouri. Insecta Mundi 7, 223-252.

Mas H, Hernández R, Villaroya MG, Sánchez G, Pérez-Laorga E, González EG, Ortiz AL, Lencina J, Rovira J, Marco M, Pérez, Gil MAI, Sánchez-García FJ, Bordón P & Pastor C (2013) Comportamiento de dispersión y capacidad de vuelo a larga distancia de Monochamus galloprovincialis (Olivier 1795). 6th Congress of Forestry, Spain.

Miller DR, Dodds KJ, Eglitis A, Fettig CJ, Hofstetter RW, Langor DW, Mayfield AE, Munson AS, Poland TM & Raffa KF (2013) Trap lure blend of pine volatiles and bark beetle pheromones for Monochamus spp. (Coleoptera: Cerambycidae) in pine forests of Canada and the United States. Journal of Economic Entomology 106, 1684-1692.

Naves P, De Sousa E & Quartau JA (2006) Reproductive traits of Monochamus galloprovincialis (Coleoptera: Cerambycidae) under laboratory conditions. Bulletin of Entomological Research 96, 289-294.

Ogawa S & Hagiwara Y (1980) Expansion of the pine infestation caused by pine wood nematodes. Shinrin Boeki (Forest Pests) 29, 115–117.

Osada Y, Yamakita T, Shoda-Kagaya E, Liebhold AM & Yamanaka T (2018) Disentangling the drivers of invasion spread in a vector-borne tree disease. Journal of Animal Ecology, 1–13. https://doi.org/10.1111/1365-2656. 12884

Pajares JA, Álvarez G, Ibeas F, Gallego D, Hall DR & Farman DI (2010) Identification and field activity of a male-produced aggregation pheromone in the pine sawyer beetle, Monochamus galloprovincialis. Journal of Chemical Ecology 36, 570-583.

Pajares JA, Álvarez G, Hall DR, Douglas P, Centeno F, Ibarra N, Schroeder M, Teale SA, Wang Z, Yan S, Millar JG & Hanks LM (2013) 2-(Undecyloxy)-ethanol is a major component of the male‐produced aggregation pheromone of Monochamus sutor. Entomologia Experimentalis et Applicata 149, 118-127.

Pershing JC & Linit MJ (1986a) Development and seasonal occurrence of Monochamus carolinensis (Coleoptera: Cerambycidae) in Missouri. Environmental Entomology 15, 251-253.

Pershing JC & Linit MJ (1986b) Biology of Monochamus carolinensis (Coleoptera: Cerambycidae) on Scotch pine in Missouri. Journal of the Kansas Entomological Society 59, 706-711.

Rasatti,D, Toffolo EP Battisti A & Faccoli M (2012) Monitoring of the pine sawyer beetle Monochamus galloprovincialis by pheromone traps in Italy. Phytoparasitica 40, 329–336.

Robinet C, Roques A, Pan H, Fang G, Ye J, Zhang Y & Sun J (2009) Role of human-mediated dispersal in the spread of the pinewood nematode in China. PLoS One 4, e4646.

Rose AH (1957) Some notes on the biology of Monochamus scutellatus (Say) (Coleoptera: Cerambycidae). Canadian Entomologist 89, 547-553.

Ryall K, Silk P, Webster RP, Gutowski JM, Meng Q, Li Y, Gao W, Fidgen J, Kimoto T, Scarr T, Mastro V & Sweeney JD (2015) Further evidence that monochamol is attractive to Monochamus (Coleoptera: Cerambycidae) species, with attraction synergised by host plant volatiles and bark beetle (Coleoptera: Curculionidae) pheromones. Canadian Entomologist 147, 564-579.

Shimazu M, Tsuchiya D, Sato H & Kushida T (1995) Microbial control of Monochamus alternatus Hope (Coleoptera: Cerambycidae) by application of nonwoven fabric strips with Beauveria bassiana (Deuteromycotina: Hyphomycetes) on infested tree trunks. Applied Entomology and Zoology 30,207-213.

Shimazu M (2009) Use of Microbes for Control of Monochamus alternatus, Vector of the Invasive Pinewood Nematode. In: Hajek, A.E., Glare, T.R., O’Callaghan, M. (eds) Use of Microbes for Control and Eradication of Invasive Arthropods. Progress in Biological Control, vol 6. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-8560-4_9

Teale SA, Wickham JD, Zhang F, Su J, Chen Y, Xiao W, Hanks LM & Millar JG (2011) A male-produced aggregation pheromone of Monochamus alternatus (Coleoptera: Cerambycidae), a major vector of pine wood nematode. Journal of Economic Entomology 104, 1592–1598.

Togashi K (1997) Lifetime fecundity and body size of Monochamus alternatus (Coleoptera: Cerambycidae) at a constant temperature. Japanese Journal of Entomology 65, 458–470.

Togashi K & Magira H (1981) Age-specific survival rate and fecundity of the adult Japanese pine sawyer, Monochamus alternatus Hope (Coleoptera: Cerambycidae), at different emergence times. Applied Entomology and Zoology 16, 351–361.

Togashi K, Appleby JE, Sadeghi HO & Malek RB (2009) Age specific survival rate and fecundity of Monochamus carolinensis (Coleoptera: Cerambycidae) under field conditions. Applied Entomology and Zoology 44, 249–256.

Torres-Villa LM, Zugasti C, De-Juan JM, Oliva MJ, Montero C, Mendola FJ, Conejo Y, Sánchez Á, Fernández, F, Ponce F & Espárrago G (2015) Mark-recapture of Monochamus galloprovincialis with semiochemical-baited traps: Population density, attraction distance, flight behaviour and mass trapping efficiency. Forestry an International Journal of Forest Research 88, 224–236.

Walsh KD & Linit MJ (1984) Feeding preferences of the adult pine sawyer, Monochamus carolinensis (Coleoptera: Cerambycidae), for four pine species. Environmental Entomology 13, 1164-1166.

Walsh KD & Linit MJ (1985) Oviposition biology of the pine sawyer, Monochamus carolinensis (Coleoptera: Cerambycidae). Annals of the Entomological Society of America 78, 81-85.

Webb JL (1909) Some insects injurious to forests. The southern pine sawyer. USDA Bulletin 58, 41-56.

Wilson LF (1975) White spotted sawyer. In Forest Pest Leaflet 74. USDA, Forest Service, Northern Area State & Private Forestry.

This datasheet was prepared in 2023 by Süleyman Akbulut. His valuable contribution is gratefully acknowledged.

EPPO (2024) Monochamus carolinensis. EPPO datasheets on pests recommended for regulation. https://gd.eppo.int (accessed 2024-12-25)

This datasheet was first published online in 2023. It is maintained in an electronic format in the EPPO Global Database. The sections on 'Identity', ‘Hosts’, and 'Geographical distribution' are automatically updated from the database. For other sections, the date of last revision is indicated on the right.