Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Neodiprion sertifer
(European pine sawfly)

Toolbox

Datasheet

Neodiprion sertifer (European pine sawfly)

Summary

  • Last modified
  • 20 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Neodiprion sertifer
  • Preferred Common Name
  • European pine sawfly
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • N. sertifer is an invasive pest species, of which a good example is its introduction and spread in North America.

Don't need the entire report?

Generate a print friendly version containing only the sections you need.

Generate report

Pictures

Top of page
PictureTitleCaptionCopyright
A female and eggs of N. sertifer.
TitleFemale
CaptionA female and eggs of N. sertifer.
CopyrightMartti Varama
A female and eggs of N. sertifer.
FemaleA female and eggs of N. sertifer.Martti Varama
Eggs and first-instar larvae of N. sertifer.
TitleEggs and first-instar larvae
CaptionEggs and first-instar larvae of N. sertifer.
CopyrightMartti Varama
Eggs and first-instar larvae of N. sertifer.
Eggs and first-instar larvaeEggs and first-instar larvae of N. sertifer.Martti Varama
Feeding larval colony of N. sertifer.
TitleLarval colony
CaptionFeeding larval colony of N. sertifer.
CopyrightMartti Varama
Feeding larval colony of N. sertifer.
Larval colonyFeeding larval colony of N. sertifer.Martti Varama
Full-grown larvae of N. sertifer.
TitleMature larvae
CaptionFull-grown larvae of N. sertifer.
CopyrightMartti Varama
Full-grown larvae of N. sertifer.
Mature larvaeFull-grown larvae of N. sertifer.Martti Varama
A Scots pine stand defoliated by N. sertifer.
TitleDamage symptoms
CaptionA Scots pine stand defoliated by N. sertifer.
CopyrightMartti Varama
A Scots pine stand defoliated by N. sertifer.
Damage symptomsA Scots pine stand defoliated by N. sertifer.Martti Varama
Larvae of N. sertifer killed by the nuclear polyhedrosis virus disease (NsNPV).
TitleNatural enemy
CaptionLarvae of N. sertifer killed by the nuclear polyhedrosis virus disease (NsNPV).
CopyrightMartti Varama
Larvae of N. sertifer killed by the nuclear polyhedrosis virus disease (NsNPV).
Natural enemyLarvae of N. sertifer killed by the nuclear polyhedrosis virus disease (NsNPV).Martti Varama

Identity

Top of page

Preferred Scientific Name

  • Neodiprion sertifer (Geoffroy in Fourcroy, 1785)

Preferred Common Name

  • European pine sawfly

Other Scientific Names

  • Diprion basalis Sakisaka
  • Diprion rufum Kl.
  • Diprion rufus
  • Diprion sertifer Geoffr.
  • Lophyrus basalis Matsumura
  • Lophyrus piceae Lepeletier
  • Lophyrus rufus Latreille; Klug; Panz; Ratz.; Retz.
  • Lophyrus sertifer Enslin
  • Neodiprion rufus
  • Neodiprion sertifer Fourcroy
  • Neodiprion sertifera Yano
  • Pteronus sertifer Geoffr.
  • Tenthredo juniperi Christ
  • Tenthredo pectinata rufa Retzius
  • Tenthredo pini rufa Villers
  • Tenthredo rufa Latreille
  • Tenthredo sertifera Geoffroy

International Common Names

  • English: fox-coloured sawfly; pine, sawfly, European; pine, sawfly, lesser; sawfly, fox-coloured
  • French: diprion du pin; diprion du pin sylvestre; lophyre roux; tenthrede du pin d'ecosse
  • Russian: ryzij sosnovyj pililschik

Local Common Names

  • Czech Republic: hrebenule rysava; pilatky rysave
  • Denmark: rød fyrrehveps
  • Estonia: punakas männivaablane
  • Finland: ruskea mäntypistiäinen; ruskomäntypistiäinen
  • Germany: Buschhornblattwespe, Rote Kiefern-; Kiefernbuschhornblattwespe, Rotgelbe; Rote Kiefernbuschhornblattwespe; Rotgelbe Kiefern-Buschhornblattwespe
  • Italy: tenthredine nerastra del pino
  • Japan: Matu-no-ki-habati
  • Netherlands: Dennebladwesp, rode; rod dennenbladwesp
  • Norway: rød furubarveps
  • Poland: borecznik rudy
  • Sweden: röd tallstekel

EPPO code

  • NEODSE (Neodiprion sertifer)

Summary of Invasiveness

Top of page N. sertifer is an invasive pest species, of which a good example is its introduction and spread in North America.

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Hymenoptera
  •                         Family: Diprionidae
  •                             Genus: Neodiprion
  •                                 Species: Neodiprion sertifer

Notes on Taxonomy and Nomenclature

Top of page Members of the Diprionidae are commonly called conifer sawflies or diprionids. This is a small family with about 125 species in 11 genera (Smith, 1993; Viitasaari, 2002). According to Ross (1951), N. sertifer was first named by Geoffroy in 1785 as Tenthredo sertifera (Fourcroy, 1785). It has subsequently been included in the genera, Lophyrus, Pteronus and Diprion. The generic designation of Neodiprion dates from 1918 (Rohwer, 1918). Ross (1955) sketched the evolution of the genus based on morphological evidence. Taxonomically, N. sertifer is a distinct species. For a long time it was the only species of the genus in Eurasia, then in the 1980s many new Neodiprion species were described from China (Xiao et al., 1984, 1985). In North America, approximately 37 Neodiprion species are known, some of which form taxonomically difficult species groups and geographic races (Knerer and Atwood, 1973; Smith, 1979; McMillin and Wagner, 1993). According to Ross (1955), the origin of N. sertifer was in North America from where it (or its ancestral form) migrated via the Bering land bridge to northern Asia and then over the whole of northern Eurasia.

Description

Top of page Eggs

The eggs are elongate-oval, 1.7-1.8 mm long and 0.3 mm wide. They are white or pale yellow to nearly white (Wilson, 1971). The eggs are laid into slits or egg pockets, which the female cuts with her saw-like ovipositor into the edge of the needle (Ghent, 1959). One egg is laid per pocket and one to 19 (average six to eight) per needle. They are embedded in the needles and are not visible. The distance between the egg pockets is 1-1.75 mm, and they can be seen as a row of semi-lunar yellow spots in the edge of the needle. However, in the spring the pockets open and the swelled eggs become more exposed. The chorion of the parasitized eggs turns blackish or brownish depending on the parasitoid species (Pschorn-Walcher and Eichhorn, 1973).

Larvae

The newly hatched larva is 3.5 mm long. The male larvae have four feeding instars with shiny black head capsules and the female larvae have five. The mature larva moults to the final non-feeding prepupal or pre-spinning larva, which spins the cocoon. Sometimes both sexes may pass through an extra feeding instar (Lyons, 1964; Juutinen, 1967). The body of L1-L2 instars is a uniform green-grey and longitudinal stripes appear in L3. A mature larva is 18 to 25 mm long and striped. The skin is densely spined. The mid-dorsal surface has a light grey to whitish narrow line, flanked by a broader green-grey line on either side. Each side of the body has two very dark lines that are separated by a thin, whitish spiracular line. The dark lines may be nearly black and tend to break up into spots. A dark spot is present in the upper area on each side of the last abdominal segment. The ventral side is light grey-green. The top of the head of the pre-spinning larva is grey-brown and paler below the eyes. The body has a dark, broken double mid-dorsal line and dark quadrangular pleural markings on a greyish background (Scheidter, 1934; Wilson, 1971).

Cocoon (pupa)

The cocoon is cylindrical with bluntly rounded ends. It is light to dark golden-brown, tough and finely textured. The male cocoons are smaller (7.5-8.5 mm) than those of the females (8.5-11.0 mm). The cocoons can be sexed by size in low-density populations, but during outbreaks the male and female cocoons may overlap in size due to them starving.

Adults

The males are 6-9.5 mm long and black. The ventral part of the abdomen is reddish-brown in total or in part and sometimes the dorsal part is reddish. The legs are yellow-brown and the coxae are black. The antennae are black, bipectinate and have 24-31 segments.

The females are 7-10.5 mm long. The head and body are brown-reddish-yellow and the base of the tibia is whitish, sometimes with dark marks on the mesonotum. The eyes and pronotum are dark. The antennae have 18-28 segments, a dark flagellum and are serrate. The common names of N. sertifer in many languages refer to the colour of the female. The female of the Chinese Neodiprion dailingensis is very similar to that of N. sertifer, but differs from the latter in having the lancet with only eight annuli instead of nine (Xiao et al., 1985).

Some gynandromorphic adults of N. sertifer, exhibiting characteristics of both sexes, have been described (Watson, 1955; Heliövaara et al., 1992; Martini et al., 1999).

Distribution

Top of page N. sertifer is the most widely distributed of the diprionids. It is native to Europe and Asia and has been introduced in north-eastern North America. In Europe, it occurs from the Mediterranean to the northern parts of Fennoscandia, from lowlands to mountains up to 2100 m. In Asia, it occurs in Siberia, Korea and Japan, even at a height of 3000 m (Pschorn-Walcher, 1965, 1982).

N. sertifer was probably introduced from Europe to North America. It was first recorded in New Jersey, USA in 1925 and in Ontario, Canada in 1939. It rapidly dispersed and was established as an important pest in south-eastern Canada and in north-eastern USA (Lyons, 1964; Griffiths et al., 1984).

Distribution Table

Top of page

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

ArmeniaPresentNative Not invasive Kolomiets et al., 1972
Georgia (Republic of)Present, few occurrencesNative Not invasive Kolomiets et al., 1972
JapanPresentPresent based on regional distribution.
-HonshuWidespreadNative Not invasive Takeuchi, 1940; Nakamura, 1982; Morimoto and Nakamura, 1989
KazakhstanWidespreadNative Not invasive Gninenko, 1995; Kolomiets et al., 1972
Korea, DPRPresentNative Not invasive Takeuchi, 1940
Korea, Republic ofPresentNative Not invasive Takeuchi, 1940
TurkeyRestricted distributionNative Not invasive cuhadar et al., 2000; Yaman et al., 2001

North America

CanadaRestricted distributionIntroduced1939 Invasive Lyons, 1964; Griffiths et al., 1984
-Newfoundland and LabradorRestricted distributionIntroduced1974 Invasive Clark and Singh, 1975; Griffiths et al., 1984; Hall, 1996
-Nova ScotiaRestricted distributionIntroduced1980 Invasive Magasi, 1981; Griffiths et al., 1984; Hall, 1996
-OntarioWidespreadIntroduced1939 Invasive Brown, 1940; Raizenne, 1957; Lyons, 1964; Griffiths et al., 1984
-QuebecWidespreadIntroduced1974 Invasive Martineau & Lavallée, 1975; Griffiths et al., 1984
USARestricted distributionIntroduced1925 Invasive Schaffner, 1939; Lyons, 1964; Coppel and Benjamin, 1965; Wilson, 1971
-ConnecticutRestricted distributionIntroduced Invasive Lyons, 1964
-DelawareRestricted distributionIntroduced1979 Invasive USDA, 1980
-IllinoisRestricted distributionIntroduced Invasive Benjamin et al., 1955; Lyons, 1964
-IndianaRestricted distributionIntroduced Invasive Lyons, 1964
-IowaRestricted distributionIntroduced Invasive Lyons, 1964; Smith, 1979
-KentuckyPresentRieske et al., 2001
-MaineRestricted distributionIntroduced Invasive Smith, 1979
-MichiganRestricted distributionIntroduced Invasive Lyons, 1964
-MissouriRestricted distributionIntroduced Invasive Lyons, 1964
-MontanaRestricted distributionIntroduced Invasive Smith, 1979
-New JerseyWidespreadIntroduced1925 Invasive Schaffner, 1939; Lyons, 1964; Smith, 1979
-New YorkRestricted distributionIntroduced Invasive Lyons, 1964
-OhioRestricted distributionIntroduced Invasive Anon., 1959
-PennsylvaniaRestricted distributionIntroduced Invasive Lyons, 1964
-South DakotaRestricted distributionIntroduced Invasive Lyons and, 1977; Smith, 1979
-WisconsinRestricted distributionIntroduced Invasive Lyons, 1964

Europe

AustriaWidespreadNative Not invasive Niklas and Franz, 1957; Jahn and Sinreich, 1964; Pschorn-Walcher, 1965; Liston, 1995; Krehan, 1999
BelarusWidespreadNative Not invasive Kharitonova & Zukhov, 1989; Rywkin, 1957
BelgiumRestricted distributionNative Not invasive Breny, 1957; Niklas and Franz, 1957
BulgariaWidespreadNative Not invasive Daskalova, 1970; Tsankov et al., 1980
CroatiaRestricted distributionNative Not invasive Opalicki, 1980; Glavas et al., 1999
Czech RepublicWidespreadNative Not invasive Simandl, 1993; Simandl and Anderbrant, 1995
Czechoslovakia (former)WidespreadNative Not invasive Martinek, 1972; Martinek, 1974; Simandl, 1989
DenmarkRestricted distributionNative Not invasive Borries, 1895; Bejer, 1989
EstoniaWidespreadNative Not invasive Kuusik and Kopvillem, 1970; Mihkelson, 1977; Mihkelson, 1980; Voolma, 2000
FinlandWidespreadNative Not invasive Kangas, 1941; Kangas, 1963; Juutinen, 1967; Viitasaari and Varama, 1987
FranceWidespreadNative Not invasive Joly, 1953; Delplanque et al., 1987
GermanyWidespreadNative Not invasive Escherich, 1942; Niklas and Franz, 1957
GreeceRestricted distributionNative Not invasive Avtzis, 1989; Johansson et al., 2001
HungaryRestricted distributionNative Not invasive Csóka, 1998; Kolonits, 1965; Dulinafka et al., 1983; Liston, 1995
IrelandPresentNative Not invasive Pschorn-Walcher, 1965
ItalyRestricted distributionNative Not invasive Pollini, 1979; Baronio et al., 1989; Baronio et al., 1997
LatviaRestricted distributionNative Not invasive Eglite and Zarins, 1993; Smits et al., 1996
LithuaniaRestricted distributionNative Not invasive Niklas and Franz, 1957; Ziogas and Zolubas, 1998; Zolubas, 1999
LuxembourgIndigenous, localizedNative Not invasive Liston, 1995
MacedoniaRestricted distributionNative Not invasive Kusevska, 1973
NetherlandsRestricted distributionNative Not invasive Hein, 1956
NorwayWidespreadNative Not invasive AustarÕ et al., 1987; AustarÕ, 1969; AustarÕ, 1971
PolandWidespreadNative Not invasive Sitowski, 1925; Gornas, 1968; Kolk and Sierpinski, 1999
PortugalPresentNative Not invasive Cardoso Cabral & Morais Figo, 1965
RomaniaRestricted distributionNative Not invasive Constantineanu, 1983; Mihalache et al., 1998
Russian FederationWidespreadNative Not invasive Kolomiets et al., 1972
-Central RussiaWidespreadNative Not invasive Kolomiets et al., 1972
-Eastern SiberiaPresentNative Not invasive Kolomiets et al., 1972
-Northern RussiaWidespreadNative Not invasive Kolomiets et al., 1972
-Russian Far EastPresentNative Not invasive Kolomiets et al., 1972
-Southern RussiaWidespreadNative Not invasive Kolomiets et al., 1972
-Western SiberiaWidespreadNative Not invasive Kolomiets et al., 1972
SlovakiaRestricted distributionNative Not invasive Leontovyc et al., 1999
SloveniaRestricted distributionNative Not invasive Jurc, 2001
SpainRestricted distributionNative Not invasive Schönwiese, 1935; Garcia-Viedma and Robredo-Junco, 1962
SwedenWidespreadNative Not invasive Forsslund, 1945; Olofsson, 1987
SwitzerlandRestricted distributionNative Not invasive Niklas and Franz, 1957; Pschorn-Walcher, 1965
UKRestricted distributionNative Not invasive Crooke, 1957; Rivers and Crooke, 1962; Britton, 1985; Liston, 1995
-ScotlandPresentTrewhella et al., 2000
UkraineWidespreadNative Not invasive Meshkova, 1998; Meshkova, 1999
Yugoslavia (former)Restricted distributionNative Not invasive Zivojinovic, 1967; Zivojinovic and Sidor, 1975

Habitat List

Top of page
CategorySub-CategoryHabitatPresenceStatus
Terrestrial

Hosts/Species Affected

Top of page N. sertifer attacks most species of the two-needled pines, but also some soft pines. In Europe, the main host trees are Pinus sylvestris, Pinus nigra, Pinus montana [Pinus mugo] and Pinus cembra. Heavy feeding has also been found on exotic pines such as Pinus strobus, Pinus banksiana, Pinus resinosa, Pinus radiata and some Japanese pines (Pschorn-Walcher, 1965, 1982). The Canadian Pinus contorta is sometimes more heavily attacked than P. sylvestris (Rivers and Crooke, 1962; Delplanque et al., 1987; Olofsson, 1988b, 1989).

In Siberia, N. sertifer occurs on Pinus sibirica (Kolomiets et al., 1972). In Japan, the host plants in the lowlands are Pinus densiflora, Pinus thunbergii and P. resinosa, and in the mountains, Pinus pumila (Suzuki, 1964; Nakamura, 1982; Morimoto and Nakamura, 1989).

In North America, the favoured hosts are the exotic P. sylvestris, P. densiflora and P. mugo, and the native P. resinosa, P. banksiana, Pinus ponderosa, P. radiata, Pinus echinata, Pinus rigida, P. strobus and many others (Craighead, 1950; Lyons, 1964). Because the range of hosts that are accepted in laboratory feeding tests is often greater than that observed in the field, it is assumed that the female oviposits on a narrower range of species than are potentially suitable for larval growth (Heitland and Pschorn-Walcher, 1993). Pinus peuce is unsuitable, and P. strobus is unfavourable for oviposition. The eggs are lost on Pinus palustris and on P. rigida when the needles die and drop (Benjamin et al., 1955; Wilson, 1971). According to Lyons (1964), no oviposition has been seen in the field on soft pines in Europe or North America. The susceptibility of a host tree to pest attack may also be dependent on the provenance or variety (Wright et al., 1967; Olofsson, 1989; Eliason and McCullough, 1997; Trewhella et al., 1997, 2000). Occasional feeding on spruces (Picea spp.) has been recorded where these grow in close proximity to pines (Gäbler, 1940; Forsslund, 1945; Crooke, 1957; Rivers and Crooke, 1962).

Growth Stages

Top of page Vegetative growing stage

Symptoms

Top of page Large larval colonies feed on needles from spring to mid-summer. The older needles are fed on down to the needle sheath and the newly developing needles are mainly untouched. The older larvae may also consume bark from the older, thin branches. Heavy outbreaks may result in the complete removal of the old foliage. Normally the trees recover later in the summer when the new shoots and needles reach their full size. Lyons (1964) has published illustrative photos of the stages in the defoliation of one tree. Heavy defoliations in subsequent years may kill buds and twigs. Although extensive mortality seldom occurs, repeated defoliation weakens the trees and increases their susceptibility to attack from secondary pests (e.g. bark beetles).

The frass of diprionid larvae can be easily distinguished from lepidopteran frass (Escherich, 1942). The shape of the diprionid excrement pellets is rhomboid. They are initially green and later turn brownish. During heavy larval infestations, a light patter caused by the dropping frass can be heard in quiet forests and a thin layer of frass can be detected on the ground.

List of Symptoms/Signs

Top of page
SignLife StagesType
Leaves / external feeding
Leaves / frass visible
Stems / external feeding
Stems / visible frass
Whole plant / plant dead; dieback

Biology and Ecology

Top of page The vast amount of literature from around the world on the biology and ecology of N. sertifer has been reviewed by Escherich (1942), Niklas and Franz (1957), Lyons (1964), Kolomiets et al. (1972), Martinek (1972, 1974), Pschorn-Walcher (1965, 1982) and Baronio et al. (1997). Wagner and Raffa (1993) have dealt with many aspects of the sawfly's life history.

N. sertifer is univoltine. The life cycle is fairly uniform throughout the whole distribution area, although temporal differences exist depending on latitude and altitude. The adults emerge from cocoons in the autumn, from August to November. The sex ratio is mostly female-biased (Craig and Mopper, 1993). The female deposits her eggs in the needles of a current year shoot and an egg cluster is formed. The number of eggs per female ranges from 30 to 140 eggs. N. sertifer is arrhenotokous i.e. unfertilized eggs produce haploid males.

Embryonic development begins in the autumn but is suspended during the winter. According to Niklas and Franz (1957), development continues throughout the winter and is retarded only by low temperatures. Breny (1956) considers the cessation as a pseudo-diapause resulting from dehydration and low availability of water from the needle tissues. According to Wallace and Sullivan (1973), a true diapause is involved. The overwintering eggs are cold-hardy and average freezing points ranging from -30°C to -37°C have been recorded (Sullivan, 1965; Juutinen, 1967; Kuusik and Kopvillem, 1970, Austarå, 1971).

The larvae hatch in the Spring, from April to the beginning of June. They feed gregariously on 1 year or older needles, leaving the new growing foliage mainly untouched. From June to July the non-feeding, pre-spinning larvae spin cocoons in the litter and humus layer in the ground and only occasionally in the foliage. The eonymphs in cocoons enter a Summer diapause, which is longest in the southern and lowland populations and shortest in the northern and alpine populations. Also, insects that spin cocoons early in the season undergo a longer eonymphal diapause than those spinning later (Lyons and Griffiths, 1962). The diapause may last for 1 year or more. The incidence of prolonged diapause increases with both latitude and altitude in Europe. It is 0-10% in the southern lowland areas and 50-100% in the northern countries (Juutinen, 1967; Mihkelson, 1977; Pschorn-Walcher, 1982). In the high mountains and in the north, a life cycle of 2 years is usual (Okutani and Ito, 1957; Austarå, 1969; Pschorn-Walcher, 1970; Martinek, 1972).

The population density of N. sertifer fluctuates in wide ranges. Outbreaks, resulting in the heavy defoliation of pine stands, are known throughout the whole distribution area. The outbreaks rise to a peak in 1 to 3 years, and collapse abruptly. Niklas and Franz (1957), Martinek (1972) and Pschorn-Walcher (1965, 1982) provided lists of the outbreaks recorded in Europe.

The outbreaks in Europe tend to occur synchronously over large areas at intervals of 6 to 13 years. The extent of the outbreaks varies considerably, from a few to 200 000 ha. The young pine stands on poor soils and with low water tables are most affected. Also the mature pine stands in Fennoscandia are frequently attacked.

In North America, most outbreaks have been unsynchronized, independent events initiated by the discovery of young, previously uninfested pine forests containing few natural control agents. Also the population levels have reached higher levels than in Europe (Lyons, 1964, 1977).

There seems little doubt that the inception of outbreaks is climatically regulated, either directly or indirectly, and that dry, hot weather is the critical factor. The factors responsible for the decline of an outbreak may vary in their effect from place to place. The most frequently recorded regulators are viral disease, parasitoids and predators.

There are four major hypotheses that tend to explain patterns of sawfly dynamics. The regulatory factors are microparasites (virus), parasitoids, generalist cocoon predators and plant defence mechanisms (Hanski, 1987). The dynamics of N. sertifer can be best explained by cocoon predation by shrews and small omnivorous mammals (Hanski, 1987, 1990; Larsson et al., 1993).

Natural enemies

Top of page
Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Adelognathus marginellus Parasite Larvae
Agonum ericeti Predator
Agrothereutes adustus Parasite Pupae
Agrothereutes adustus
Aranea ventricosa Predator China
Bacillus thuringiensis Pathogen
Bacillus thuringiensis kurstaki Pathogen
Bacillus thuringiensis thuringiensis Pathogen
Beauveria bassiana Pathogen
Birdiavirus diprionis Pathogen
Blondelia inclusa Parasite Larvae
Blondelia nigripes Parasite Larvae
Clethrionomys glareolus Predator Pupae
Closterocerus ovularum Parasite Canada Pinus
Closterocerus ruforum Parasite Eggs Canada Pinus
Dahlbominus fuscipennis Parasite Pupae Canada; USA Pinus
Delomerista diprionis Parasite Pupae
Diplostichus hamatus Parasite Larvae
Dipriocampe diprioni Parasite Eggs Canada Pinus
Drino angustifacies Parasite Larvae
Drino bohemica Parasite Larvae
Drino gilva Parasite Larvae
Drino inconspicua Parasite Larvae
Endasys subclavatus Parasite Pupae
Eulophus pennicornis Parasite
Exenterus abruptorius Parasite Larvae Canada; USA Pinus
Exenterus adspersus Parasite Larvae
Exenterus affinis Parasite Larvae
Exenterus amictorius Parasite Larvae
Exenterus nigrifrons Parasite Larvae
Formica cinerea Predator
Formica exsectoides Predator
Formica polyctena Predator
Lamachus eques Parasite Larvae Canada; USA Pinus
Lophyroplectus luteator Parasite Larvae Canada; Ontario; USA Pinus
Macroneura vesicularis Parasite Pupae
Mastrus aciculatus Parasite
Mastrus argeae Parasite Pupae
Misumenops tricuspidatus Predator China
Neochrysocharis formosa Parasite Eggs
Nucleopolyhedrosis virus Pathogen Larvae
Ophion abbreviator Parasite Canada Pinus
Oxyopes ramosus Predator
Paecilomyces farinosus Pathogen
Pardosa hyperborea Predator
Pardosa sphagnicola Predator
Parus major Predator Larvae
Pleolophus basizonus Parasite Pupae Newfoundland; Ontario; USA Pinus
Pleolophus indistinctus Parasite Pupae
Prosternon tessellatum Predator
Pterostichus nigrita Predator
Sorex caecutiens Predator
Synomelix scutulata Parasite Larvae
Trineptis Parasite Pupae
Vibrissina spinigera Parasite Larvae
Villa sinuosa Parasite Pupae
Xysticus audax Predator

Notes on Natural Enemies

Top of page N. sertifer is attacked by several hymenopterous and dipterous parasitoids, and many predators including ants, bugs, beetles, lacewings, spiders, small mammals and birds. Pathogenic fungi, bacteria and a species-specific nuclear polyhedrosis virus (NsNPV) also attack it.

The diprionid larvae have special defence strategies against predators and parasitoids (Codella and Raffa, 1993) and being gregarious is one of these. The larvae also exhibit various alarm reactions and defensive displays. Prop (1960) distinguished three types of larval displays: U-bend; jerking; and stretching. The larvae possess a pair of oesophageal diverticulae with resinous contents. In response to harassment, the larvae regurgitate a droplet of the resinous liquid. This behaviour has been shown to repel ants, pentatomid bugs, spiders and birds, as well as hymenopteran and dipteran parasitoids.

There are a large number of parasitoid records from N. sertifer in the literature. However, according to Oehlke (1965) and Pschorn-Walcher (1965), a relatively high proportion of these records, particularly the older ones, are unreliable. The main sources of error have been: the mass-collection and mass-rearing of larvae and cocoons; the inadequate sample-size and faulty timing of sampling; and the false identification of the parasitoids or the hosts. The list of parasitoids presented here is based mainly on the reviews and articles of Niklas and Franz (1957), Rywkin (1957), Finlayson and Finlayson (1958), Finlayson (1960), Lyons (1964), Oehlke (1965), Kolomiets et al. (1972), Martinek (1972, 1974), and Pschorn-Walcher (1965, 1967, 1982).

Four chalcidoid egg parasitoids are known from N. sertifer in Europe (Pschorn-Walcher and Eichhorn, 1973; Pschorn-Walcher, 1982). The tetracampid Dipriocampe diprioni is strictly primary, the eulophids Closterocerus ruforum and C. formosus act as primary and hyperparasitoids, even as autoparasitoids, while the eulophid Baryscapus (Tetrastichus) oophagus is a rare hyperparasitoid. As primary egg parasitoids, D. diprioni and C. ruforum are the dominant species. In Europe, egg parasitism (including host-feeding) is generally between 5 and 50%, but during the break-down phase of the host population, much higher values have been recorded, such as from 90-95% (Juutinen, 1967; Pschorn-Walcher and Eichhorn, 1973; Juutinen and Varama, 1986). The egg parasitoids of N. sertifer do not occur in North America, and attempts to introduce and colonize them from Europe have been unsuccessful (McGugan and Coppel, 1962; Griffiths et al., 1984).

The most important parasitoids of N. sertifer in Europe are the larval parasitoids, Lamachus eques, Lophyroplectus luteator, Exenterus abruptorius, and the cocoon parasitoids Pleolophus basizonus (Ichneumonidae) and Dahlbominus fuscipennis (Eulophidae). Of minor importance are Exenterus amictorius, Exenterus adspersus, Synomelix scutulata (Ichneumonidae), and Drino inconspicua and Blondelia inclusa (Tachinidae) as larval parasitoids, and the ichneumonid, Agrothereutes adustus as a cocoon parasitoid (Pschorn-Walcher, 1965, 1967, 1982).

The parasitoid complex of N. sertifer in North America consists of about 30 native primary parasitoids, some native hyperparasitoids, and six introduced and established European parasitoids (Lyons 1964; Griffiths et al., 1984). The important native species are: the larval parasitoids, Exenterus nigrifrons [Exenterus canadensis], Exenterus affinis (Ichneumonidae), Spathimeigenia spinigera [Vibrissina spinigera], Diplostichus hamatus (Tachinidae); and the cocoon parasitoids, Pleolophus indistinctus, Endasys subclavatus, Delomerista diprionis, Mastrus argeae (Ichneumonidae), Tritneptis spp. (Pteromalidae), Eupelmus vesicularis [Eupelmus vesicularis] (Eupelmidae) and Hemipenthes sinuosa [Villa sinuosa] (Bombyliidae). The established European species are the ichneumonids, E. amictorius, E. abruptorius, P. basizonus, L. luteator and the eulophid, D. fuscipennis, and the tachinid, Drino bohemica (Griffiths et al., 1984). Also the introduced torymid, Monodontomerus dentipes, has been encountered a couple of times (Finlayson, 1960; Lyons 1964).

Mortality of N. sertifer due to parasitism varies greatly (10-90%) by year and locality. The parasitoids, along with the nuclear polyhedrosis virus, play an important role especially during the break-down phase of the outbreak. They often cause the abrupt collapse of the host population (Niklas and Franz, 1957; Lyons, 1964; Pschorn-Walcher, 1965, 1982, 1988).

True hyperparasitoids do not play a very significant role among the parasitoids, but multi-parasitism in the form of competition between larval and cocoon parasitoids may be increased when the total parasitism is high (Pschorn-Walcher, 1973).

The literature on predation upon N. sertifer has been reviewed by Niklas and Franz (1957), Lyons (1964), Kolomiets et al. (1972), Martinek (1972) and Pschorn-Walcher (1965, 1982). Tits (Parus spp.), lacewings (Chrysopa ventralis), and pentatomid, reduviid and nabid bugs (Picromerus bidens, Euschistus tristigmatus, Podisus spp., Rhinocoris annulatus [Rhynocoris annulatus], Zelus sp. and Nabis sp.) have been reported to occasionally prey on the eggs of N. sertifer.

Due to the effective defence mechanisms and the unpalatability of the larvae to predators, mortality of the larval stages caused by the predators is generally low. Birds, ants (Formica spp.), predatory bugs, coccinellids, neuropterans and spiders have been reported to occasionally feed on the larvae. However, Schwerdtfeger (1936) reported that ants of the Formica rufa-group were one of the principal factors causing sawfly mortality in the Prussian plain. According to Bruns and Schrader (1955), no cocoons with living N. sertifer could be found within 30 m of F. rufa nests. In Sweden, Olofsson (1992) reported a high mortality of the larvae within 40 m of a Formica polyctena nest.

Cocoon predation plays an important role in the population dynamics of N. sertifer and of the other diprionid species. The larvae of the family Elateridae (Coleoptera) destroy cocoons in the ground, and birds (mainly Parus spp.) prey effectively on the cocoons occasionally spun in the foliage. However, the most effective cocoon predators are small mammals such as shrews, mice and voles (Soricidae, Microtidae). Holling (1959), Hanski and Parviainen (1985) and Hanski (1987, 1990) studied small mammal predation of N. sertifer. According to Hanski (1990), small mammals frequently consume 80% or more of the cocoons in endemic sawfly populations when the density of cocoons in coniferous forests in Fennoscandia is less than 10 000 per ha. In contrast, during severe outbreaks when there are several million cocoons per ha, only a small fraction is destroyed by small mammals.

A lethal species-specific nuclear polyhedrosis virus disease (NsNPV) frequently infects the larvae of N. sertifer. The disease is caused by Borrelinavirus diprionis [NeseNPV], which belongs to the baculoviridae (Cunningham and Entwistle, 1981). It occurs widely in nature and is often one of the main factors causing the abrupt collapse of outbreaks (Lyons, 1964; Pschorn-Walcher, 1982). The larvae become infected either by ingesting the polyhedral inclusion bodies (PIB) with food, or via an infected parent. The parasitoids, predators and scavengers may act as transmission agents because the virus stays infectious in their faeces (Olofsson, 1988a). NsNPV has been widely used in the biological control of N. sertifer.

Means of Movement and Dispersal

Top of page Natural dispersal (non-biotic)

N. sertifer larvae normally pupate (spin cocoons) in the litter and soil, and occasionally in the foliage or branches. During outbreaks there is a risk that the larvae pupate in forests in containers, vehicles, camping equipment etc., and thus get transported into new areas.

Silvicultural practices

N. sertifer is an introduced, dispersing species in North America. Some new infestations did not result from natural dispersal but from the introduction of infested nursery stock (Griffiths et al., 1971).

Plant Trade

Top of page
Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bark pupae Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants pupae Yes Pest or symptoms usually visible to the naked eye
Leaves eggs; larvae; pupae Yes Pest or symptoms usually visible to the naked eye
Stems (above ground)/Shoots/Trunks/Branches larvae; pupae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Roots
Seedlings/Micropropagated plants
True seeds (inc. grain)
Wood

Wood Packaging

Top of page
Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
Solid wood packing material with bark Yes
Solid wood packing material without bark Yes
Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood

Impact

Top of page N. sertifer is a univoltine, early season defoliator. The larvae feed on older needles and leave the new growing needles mainly untouched. Hence tree mortality is normally modest, even after heavy infestations. In North American Christmas tree plantations, N. sertifer has been a serious problem and control applications have been widely used, because even a light defoliation renders the trees unmarketable (Lyons, 1964; Wilson, 1971).

Relatively little is known about the effect of defoliation on tree growth. In Michigan, 20, 65, 85, and 100% defoliation before new foliage developed, caused 14, 23, 37, and 63% loss in terminal shoot elongation and 18, 47, 53, and 71% reduction in radial increment, respectively. The 100% defoliation was simulated and trees survived 3 years of complete defoliation (Wilson, 1966).

In Hungary, heavy defoliation caused 30-45% reduction in annual growth (Kolonits, 1965). In Sweden, Forsslund (1945) found the height growth loss to vary between 25% and 60% in young Scots pine stands defoliated for two consecutive years. Eklund (1964) recorded a reduction of 52% in the diameter growth of old pine stands due to defoliation. The measurements were made 1-2 years after the infestation, therefore they do not cover the whole recovery period, which may be as long as 9 years (Austarå et al., 1987).

In Finland, the mortality of Scots pine after 1 years defoliation was recorded as approximately 4%, and a reduction in the volume increment for the following 5 years was 20% corresponding to one normal annual increment (Juutinen, 1967; Tiihonen, 1970). Most of the killed trees were suppressed or weakened before the infestation. In Norway, a heavy defoliation of Scots pine for 2 years in 90-120-year-old forests caused a volume loss of 33% during 9 years. This loss corresponds to three normal annual increments (Austarå et al., 1987). The economic consequences of the growth loss and mortality have been calculated by Austarå et al. (1987), and Lyytikäinen-Saarenmaa and Tomppo (2002).

Detection and Inspection

Top of page The overwintering egg clusters, which appear as rows of yellow spots (egg pockets) in the edges of the needles in the current year's shoots, can be detected from the Autumn to the Spring by careful inspection. The signs of defoliation reveal the feeding larval colonies. The presence of cocoons in the soil can be checked. Pheromone traps containing the female sexual pheromone can be set to catch male insects at the time of their emergence from the cocoons in Autumn.

Similarities to Other Species/Conditions

Top of page The way of feeding and the symptoms of defoliation are very similar between many pine sawfly species. In Europe, N. sertifer is the only diprionid species that overwinters in the egg-stage and defoliates pines in the Spring or early Summer. For proper identification of the pest species, samples of the egg-bearing needles, larvae, larval skins, adults or cocoons should be taken.

Prevention and Control

Top of page

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.

Biological control

One of the best examples of the successful development of biological control methods in forest protection is the nuclear polyhedrosis virus of N. sertifer (NsNPV). The virus was discovered in Germany (Escherich, 1913). Bird (1953, 1955, 1961) developed a supply of the virus from a few dead larvae of N. sertifer from Sweden in 1949. This was used for experimental purposes and eventually for release as a control agent. Due to the intensive studies in America and Europe (e.g. Bird and Whalen, 1953; Franz and Niklas, 1954; Benjamin et al., 1955; Krieg, 1955; Rivers and Crooke, 1962; Nuorteva, 1972; Donaubauer, 1973; Cunningham and Entwistle, 1981; Juutinen, 1982; Podgwaite et al., 1984), the NsNPV became the most widely used baculovirus in several countries (Pschorn-Walcher, 1982; Morris et al., 1986). It has been registered under different names in the USA, UK, Finland and the Czech Republic. In Ontario, N. sertifer was a major pest of Christmas tree plantations but since the 1970s it has only been a minor pest due to the use of the NsNPV and parasitoid introductions (Griffiths et al., 1984; Morris et al., 1986).

The incubation time of NsNPV is 1-2 weeks. The virus should be applied on early instars (L1, L2) to be effective and cause over 90% mortality. According to the critical review by Olofsson (1988a), the virus plays a minor role in the population dynamics of N. sertifer; often a smaller role than other biotic mortality factors. Therefore, virus treatment should be resorted to in exceptional cases only, when there are specific indications that a treatment is required. For example, when severe damage by secondary forest insects, e.g. bark beetles, can be expected. See van Frankenhuyzen (2002) for an assesssment of the successful control of N. sertifer in Canada with the NPV.

In the laboratory and field tests performed by Heimpel and Angus (1963), Bacillus cereus was pathogenic (25% mortality) to larvae of N. sertifer, but the Bacillus thuringiensis var. sotto Ishiwata toxin was ineffective (Angus, 1956). Sezen et al. (2001) investigated the insecticidal potential of the bacterium, Serratia marcescens Bn10, which was isolated from Balaninus nucum. Mortality of the N. sertifer larvae was 88% within 5 days.

The rhabditid nematode, Heterorhabditis heliothidis, caused 100% mortality of N. sertifer larvae within 120 hours at 15°C in the laboratory. The intact cocoons were not susceptible and only 11% mortality in 168 hours was observed. However, the pupae that were dissected out of the cocoons were all parasitized in 48 hours (Finney and Bennett, 1983).

In small ornamental trees or in pine plantations of small acreage, N. sertifer can be controlled simply by hand-picking and destroying the larval colonies. A mere knocking down of the larvae is useless, because the larvae climb back into the trees within a few hours (Teräs, 1982).

Host-Plant Resistance

Different pine species, provenances and varieties have been shown to vary in their susceptibility to N. sertifer (refer to Notes on host range). Where possible, less susceptible alternatives should be used in forestry programs.

The larvae of N. sertifer normally reject the young foliage of the Scots pines. This feeding pattern is mainly caused by a deterrent compound identified as 13-keto-8(14)-podocarpen-18-oic acid (Niemelä et al., 1982). This substance was mainly found in the young foliage of pines, and the highest concentrations were found in the trees whose needles had been badly damaged by the N. sertifer larvae.

Pheromonal control

Like many other species of diprionids, the males of N. sertifer are strongly attracted by the female sex pheromones. The investigation of the diprionid sex pheromones started in the late 1950s (Coppel et al., 1960), and the first pheromone was identified 16 years later (Jewett et al., 1976). Anderbrant (1993, 1999) thoroughly reviewed the literature since then and up to 1999.

The inactive precursor of the N. sertifer pheromone is 3,7-dimethyl-2-pentadecanol (diprionol) and the active compound is (2S,3S,7S)-diprionyl acetate or propionate, which has proved to be attractive in Europe, North America and Japan (Anderbrant, 1993, 1999). Traps baited with synthetic pheromone can be used for many purposes. The flight period can be determined (Simandl, 1993; Schedl, 1994), the abiotic factors important for flight can be investigated (Jönsson and Anderbrant, 1993), and the behaviour of the sawflies can be studied (Östrand, 2001). The first attempts to use pheromone monitoring traps for determining population densities or population trends of N. sertifer showed either significant relationships (Baldassari et al., 2000), only weak relationships (Lyytikäinen-Saarenmaa et al., 1999, 2001) or no significant correlation (Herz et al., 2000) between the trap catch and the sawfly density or defoliation.

The use of sex pheromones in the mating disruption technique to control populations resulted in the suppression of the N. sertifer population in isolated pine stands in Italy (Martini et al., 2002). It was ineffective in larger areas of pine forests in Sweden (Anderbrant et al., 1995).

Chemical control

The diprionid sawflies are susceptible to most stomach and contact poisons, and a vast array of insecticides have been utilized in abatement programmes (Lyons, 1964; Coppel and Benjamin, 1965, Pschorn-Walcher, 1982). Early control programmes mainly utilized lead or calcium arsenate (Hamilton, 1943), a hazardous chemical that is not recommended.

Currently chlorinated hydrocarbons have been replaced with organophosphorous insecticides and pyrethroids, which have proved to be effective against the diprionid larvae (Wallner, 1968; Malinowski, 1995; Safronov, 1996). In North America, malathion has been used to control N. sertifer (Wilson, 1971).

The insect growth regulators, acylurea insecticides, have been actively developed and tested since the 1970s. They interfere with the normal formation of the cuticle and the deposition of chitin, and are environmentally safer than many early pesticides. Skatulla (1975) reported that aerial spraying of diflubenzuron caused 100% mortality of the N. sertifer larvae in 17 days, and no adverse side effects were observed. Ultra low volume aerial application of diflubenzuron resulted in 98-100% mortality of the N. sertifer larvae (Adomas, 1999). Malinowski (1995) demonstrated that flufenoxuron, novaluron and teflubenzuron were more active than diflubenzuron and triflumuron against the diprionid larvae. According to Skatulla (1975), diflubenzuron remains effective for 4 months. Schwenke (1979) discussed the possible environmental side effects of diflubenzuron applications.

Azadirachtin is a natural insecticide from the neem tree (Azadirachta indica) and strongly reduced the feeding activity and body weight of N. sertifer larvae under laboratory conditions. 100% mortality was achieved. For experimental treatment in the field, azadirachtin concentrations at levels from 0.001-0.00001% were proposed (Malinowski, 2002).

According to All and Benjamin (1976), certain insect antifeedant chemicals have the potential to reduce Neodiprion larval feeding. When larvae were subjected to field sprays, feeding, development and colony integrity were often disrupted. When colonies disbanded, the solitary larvae were more vulnerable to entomophages. Thus the antifeedants could be important components in an integrated control program to maximize the effectiveness of the natural control agents.

References

Top of page

+strand F, 2001. Behaviour of pine sawflies in relation to pheromone-based pest management. Dissertation. Sweden: Dep. Ecol., Chem. Ecol. Ecotoxic., Lund University.

Adomas J, 1999. The effectiveness of Dimilin 480 SC in control of Diprioninae [Diprionidae]. Progress in Plant Protection, 39(2):524-526; 3 ref.

All JN; Benjamin DM, 1976. Potential of antifeedants to control larval feeding of selected Neodiprion sawflies (Hymenoptera: Diprionidae). Canadian Entomologist, 108(11):1137-1144

Anderbrant O, 1993. Pheromone biology of sawflies. Sawfly life history adaptations to woody plants [edited by Wagner, M. R.; Raffa, K. F.] San Diego, USA; Academic Press, Inc., 119-154

Anderbrant O, 1999. Sawflies and seed wasps. In: Hardie J, Minks AK, eds. Pheromones of non-lepidopteran insects associated with agricultural plants. Wallingford, UK: CAB International, 199-226.

Anderbrant O; L÷fqvist J; Hedenstr÷m E; H÷gberg H-E, 1995. Mating disruption of the pine sawfly Neodiprion sertifer (Hymenoptera: Diprionidae): effects on pheromone trap catch, sawfly sex ratio and density and on tree damage. In: Hain FP, Salom SM, Ravlin WF, Payne TL, Raffa KF, eds. Behavior, population dynamics, and control of forest insects: Proceedings of the IUFRO Joint Conference, Feb., 1994. Ohio, USA: The Ohio State University Press, 415-427.

Angus TA, 1956. The reaction of certain lepidopterous and hymenopterous larvae to Bacillus sotto toxin. Canadian Entomologist, 88:280-283.

Anonymous, 1959. Neodiprion sertifer (Geoffr.). Distribution maps of insects pests, Series A, Map no. 98, London, UK: Commonwealth Institute of Entomology

Austars +, 1969. Prolonged diapause in Neodiprion sertifer (Geoffr.) (Hymenoptera: Diprionidae) from Gaupne in Western Norway. Meddelelser fra det Norske Skogforsoeksvesen, 27:201-213.

Austars +, 1971. Cold-hardiness in eggs of Neodiprion sertifer (Geoffroy)(Hym., Diprionidae) under natural conditions. Norsk Entomologisk Tidsskrift, 18:45-48.

Austars +; Orlund A; Svendsrud A; Veidahl A, 1987. Growth loss and economic consequences following two years defoliation of Pinus sylvestris by the pine sawfly Neodiprion sertifer in West-Norway. Scandinavian Journal of Forest Research, 2:111-119.

Avtzis N, 1989. Die wichtigsten Schadinsekten der griechischen WSlder. Allgemeine Forst Zeitschrift, No. 4:95.

Baldassari N; Martini A; Baronio P; Rocchetta G, 2000. The behaviour of Neodiprion sertifer (Geoffroy) (Hymenoptera Diprionidae) populations in Scots pine stands of the Appennino Tosco-Romagnolo. Bollettino dell'Istituto di Entomologia `Guido Grandi' della Universita^grave~ degli Studi di Bologna, 54:91-100; 15 ref.

Baronio P; Faccioli G; Butturini A, 1989. A study on the influence of defoliation by Neodiprion sertifer (Geoffr.) (Hym. Diprionidae) on the growth of Pinus sylvestris L. in Romagna. Bollettino dell'Istituto di Entomologia "Guido Grandi" della Universita degli Studi di Bologna, 43:17-24

Baronio P; Martini A; Baldassari N, 1997. Neodiprion sertifer is a dangerous pest of Scots pine forests. Informatore Fitopatologico, 47(6):12-20; 45 ref.

Bejer B, 1989. Skovbrugets skadedyr 1988. Skoven 1:14-15.

Benjamin DM; Larson JD; Drooz AT, 1955. The European pine sawfly on the Henderson State Forest, Illinois, with notes on its biology and control. Journal of Forestry, 53:349-362.

Bird FT, 1953. The use of a virus disease in the biological control of the European pine sawfly, Neodiprion sertifer (Geoff.). Canadian Entomologist, 85:437-446.

Bird FT, 1955. Virus diseases of sawflies. Canadian Entomologist, 87:124-127.

Bird FT, 1961. Transmission of some insect viruses with particular reference to ovarial transmission and its importance in the development of epizootics. Journal of Insect Pathology, 3:352-380.

Bird FT; Whalen MM, 1953. A virus disease of the European pine sawfly, Neodiprion sertifer (Geoffr.). Canadian Entomologist, 85:433-437.

Borries H, 1895. Iagttagelser over Danske Naaletrae-Insekter. Tidsskr. Skovvaesen, 7 B:1-95.

Breny R, 1956. L¦embryon de Neodiprion sertifer Geoffr. en periode hivernale. Bull. Inst. Agron. Gembloux, 24:121-130.

Breny R, 1957. Contribution a l'Ttude de la diapause chez Neodiprion sertifer Geoffr. dans la nature. MTm. Acad. Roy. Belgique Cl. Sci., 30:1-88.

Breny R; Detroux L, 1951. Le ¦lophyre roux¦ a la Reid, en 1949. Bull. Soc. for Belg., 58:68-71.

Britton R, 1985. Entomology. Effects of the European pine sawfly on lodgepole pine. Report on Forest Research, Forestry Commisssion, 66-67

Brown AWA, 1940. Annual report of the Forest Insect Survey. Ottawa, Canada: Canadian Department of Agriculture, Division of Entomology.

Bruns H; Schrader A, 1955. Abnahme der Kokondichte der Roten Kiefernbuschhornblattwespe (Neodiprion sertifer) bei Nestern der Roten Waldamaise. Waldhygiene, 1:59-61.

Cardoso Cabral MTE; Morais Figo M de LNCBP, 1965. Neodiprion sertifer (Geoff.) Hymenoptera-Diprionidae, espTcie nova para a fauna lusitanica [Neodiprion sertifer, a species new to the fauna of Portugal]. Estud. Divulg. tTc. Dir. Ger. Serv. Flor. Aquic., 16:1-18.

Clark RC; Singh P, 1975. Newfoundland region. Canadian Forest Service, Annual Report Forest Insect Disease Survey, 1974.

Codella SG Jr; Raffa KF, 1993. Defense strategies of folivorous sawflies. In: Wagner MR, Raffa KF, eds. Sawfly life history adaptations to woody plants. San Diego, USA; Academic Press, Inc., 261-294

Constantineanu R, 1983. Ichneumonidae (Hymenoptera) new and rare to the Romanian fauna. Studii si Cercetari de Biologie, Biologie Animala, 35(2):77-81

Coppel HC; Benjamin DM, 1965. Bionomics of the nearctic pine-feeding diprionids. Annual Review of Entomology, 10:69-96.

Coppel HC; Casida JE; Dauterman WC, 1960. Evidence for a potent sex attractant in the introduced pine sawfly, Diprion similis. Annals of the Entomological Society of America, 53:510-512.

Craig TP; Mopper S, 1993. Sex ratio variation in sawflies. In: Wagner MR, Raffa KF, eds. Sawfly life history adaptations to woody plants. San Diego, USA; Academic Press, Inc., 61-92

Craighead FC, 1950. Insect enemies of eastern forests. USDA, Misc. Publ. 657.

Crooke M, 1957. A brief review of the British conifer feeding sawflies. Zeitschrift fur Angewandte Entomologie, 41:179-183.

Cs=ka G, 1998. Recent trends of damage caused by pests and pathogens in the Hungarian forests. Methodology of forest insect and disease survey in Central Europe. Proceedings, First Workshop of the IUFRO WP 7.03.10, Ustron^acute~-Jaszowiec, Poland, 21-24 April, 1998., 115-120; 2 ref.

Cuhadar AGI; Aksu Y; Babacan N, 2000. Research on Neodiprion sertifer (Geoff.) creating damage in the forest areas of Artvin, Turkey. Orman Mu^umlaut~hendislig^breve~i, 37(12):18-24; 3 ref.

Cunningham JC; Entwistle PF, 1981. Control of sawflies by Baculovirus. In: Burges HD, ed. Microbial Control of Pests and Plant Diseases 1970-1980. London, UK: Academic Press, 379-407.

Daskalova I, 1970. [Distribution of Neodiprion sertifer in Bulgaria.] Nauch. Trud. Lesotekh. Inst. (Ser. Gorsko Stop.), 18:139-143.

Delplanque A; Giovanni JP; Roques A; Agustin S, 1987. First entomological problems relating to the introduction of Pinus contorta in France. Boletin de la Estacion Central de Ecologia, 16(31):123-132

Donaubauer E, 1973. Results of polyhedral virus applications against Neodiprion sertifer Geoffr. Bulletin, Organisation Europeenne et Mediterraneenne pour la Protection des Plantes, 3(3):105-110

Dulinafka G; Darvas B; Dezsery M; Gyulai P; Mezo G; Szalai-Marzso L, 1983. Control of European pine sawfly (Neodiprion sertifer Geoffr.) with nuclear polyhedrosis virus. In: Darvas B, Toth M, Vajna L, eds. Proceedings of the International Conference on Integrated Plant Protection, 4, Selective Control Methods, 4-9 July, 1983, Budapest, Hungary: Hungarian Society of Agricultural Sciences, 7-10.

Eglite G; Zarins I, 1993. The European pine sawfly nuclear polyhedrosis virus and its application to insect pest control. Latvijas Zina^macron~tn^tail~u Akade^macron~mijas Ve^macron~stis. B Dal^tail~a, Dabaszina^macron~tnes, No. 7:73-78; 22 ref.

Eklund B, 1964. Om sterverkningarna av den r÷da tallstekelns skadeg÷relse ps tallens diametertillvSxt vid br÷sth÷jd. Norrlands Skogsv. f÷rb. Tidskr., 1964:205-218.

Eliason EA; McCullough DG, 1997. Survival and fecundity of three insects reared on four varieties of Scotch pine Christmas trees. Journal of Economic Entomology, 90(6):1598-1608; 41 ref.

Escherich E, 1942. Die Forstinsekten Mitteleuropas. Vol. V. Hymenoptera (Hautflügler) und Diptera (Zweiflügler). Berlin: Paul Parey.

Escherich K, 1913. Neues nber Polyederkrankheiten. Naturw. Z. Land- Forstw., 11, 86 pp.

Finlayson LR; Finalyson T, 1958. Parasitism of the European pine sawfly, Neodiprion sertifer (Geoff.) (Hymenoptera: Diprionidae), in southwestern Ontario. Canadian Entomologist, 90:223-225.

Finlayson T, 1960. Taxonomy of cocoons and puparia, and their contents, of Canadian parasites of Neodiprion sertifer (Geoff.) (Hymenoptera: Diprionidae). Canadian Entomologist, 92:20-47.

Finney JR; Bennett GF, 1983. The susceptibility of some sawflies (Hymenoptera: Tenthredinidae) to Heterorhabditis heliothidis (Nematoda: Heterorhabditidae) under laboratory conditions. Canadian Journal of Zoology, 61(5):1177-1180

Forsslund KH, 1945. Nsgot om r÷da tallstekelns (Diprion sertifer Geoffr.) skadeg÷relse. Medd. stat. skogsf÷rs. anst., 34:365-390.

Fourcroy AFde, 1785. Entomologia Parisiensis 2. Paris, France, 544 pp.

Franz J; Niklas OF, 1954. Feldversuche zur BekSmpfung der Roten Kiefernbuschhornblattwespe (Neodiprion sertifer (Geoffr.)) durch knnstliche Verbreitung einer Virusseuche. Nachrbl. Dtsch. Pflschutzd. Braunschweig, 6:131-134.

Garcia-Viedma M; Robredo-Junco F, 1962. Insect pests in pine plantations. II Asamblea Tecnica Forestal, Ministero de Agricultura, Madrid 1962 No. 4 (Session 8a sub session A), 1963, 918-922.

Ghent AW, 1959. Row-type oviposition in Neodiprion sawflies as exemplified by the European pine sawfly, N. sertifer (Geoff.). Canadian Journal of Zoology, 37:267-281.

Glavas M; Diminic D; Hrasovec B, 1999. Diseases and pests of pines in the coastal region of Croatia. In: Forster B, Knizek M, Grodzki W, eds. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO WP 7.03.10, April 20-23, 1999, Sion-Chateauneuf, Switzerland, 229-230.

Gninenko YuI, 1995. Ecology of the sawfly Neodiprion sertifer in the pine stands of northern Kazakhstan and the southern Trans-Urals. Lesovedenie, No. 6:56-61; 16 ref.

Gornas E, 1968. Stand und Problematik des gegenwSrtigen Auftretens der Kiefernbuschhornblattwespe in Polen. Archiv fnr Forstwesen, 17:641-652.

Griffiths KJ; Cunningham JC; Otvos IS, 1984. Neodiprion sertifer (Geoffroy), European pine sawfly (Hymenoptera: Diprionidae). In: Kelleher JS, Hulme MA, eds. Biological control programmes against insects and weeds in Canada 1969-1980. Slough, UK: Commonwealth Agricultural Bureau, 331-340.

Griffiths KJ; Rose AH; Bird FT, 1971. Neodiprion sertifer (Geoff.), European pine sawfly (Hymenoptera: Diprionidae). In: Biological control programmes against insects and weeds in Canada 1959-1968. Commonwealth Institute of Biological Control Technical Communication 4, 150-162.

GSbler H, 1940. Lophyrus rufus Retz. = sertifer Geoffr. an Bergkiefer und Fichte. Anz. SchSdlingsk., 16:22-23.

Hall JP, 1996. Forest insect and disease conditions in Canada 1994. Forest Insect and Disease Conditions in Canada, No. 1994:iv + 112 pp.; 10 pp. of ref.

Hamilton CC, 1943. The pine sawfly Neodiprion sertifer (Geoff.) and its control with concentrated lead arsenate sprays. J. Econ. Ent., 36:236-240.

Hanski I, 1987. Pine sawfly population dynamics: patterns, processes, problems. Oikos, 50(3):327-335

Hanski I, 1990. Small mammal predation and the population dynamics of Neodiprion sertifer. In: Watt AD, Leather SR, Hunter MD, Kidd NAC, eds. Population Dynamics of Forest Insects. Andover, UK: Intercept Ltd, 253-263.

Hanski I; Parviainen P, 1985. Cocoon predation by small mammals, and pine sawfly population dynamics. Oikos, 45(1):125-136

Heimpel AM; Angus TA, 1963. Diseases caused by certain sporeforming bacteria. In: Steinhaus EA, ed. Insect pathology, an advanced treatise, 2:21-73. New York, USA: Academic Press, 689 pp.

Hein G, 1956. De plaag van de rode dennenbladwesp (Diprion sertifer Geoffr.) in Nederland in de jaren 1949-1954 en een onderzoek naar in de voedselplant liggende, de eiafzetting remmende factoren. Nederl. Boschbouw Tijdschr., 28:257-265, 285-297.

Heitland W; Pschorn-Walcher H, 1993. Feeding strategies of sawflies. Sawfly life history adaptations to woody plants, 93-118; 74 ref.

Heliovaara K; Vaisanen R; Varama M; Viitasaari M, 1992. Gynandromorphic individuals of Neodiprion sertifer (Hymenoptera, Diprionidae). Entomologica Fennica, 3(3):149-153

Herz A; Heitland W; Anderbrant O; Ediund H; Hedenstr÷m E, 2000. First use of pheromones to detect phenology patterns and density relationships of pine sawflies in German forests. Agricultural and Forest Entomology, 2(2):123-129; 30 ref.

Holling CS, 1959. The components of predation as revealed by a study of small mammal predation of the European pine sawfly. Canadian Entomologist, 91:293-320.

J÷nsson P; Anderbrant O, 1993. Weather factors influencing catch of Neodiprion sertifer (Hymenoptera: Diprionidae) in pheromone traps. Environmental Entomology, 22:445-452.

Jahn E; Sinreich A, 1964. Beobachtungen zur Massenvermehrung der Schwarzk÷pfigen Kiefernbushhornblattwespe, Neodiprion sertifer Geoffr. im pannonischen Klimagebiet +sterreichs in den Jahren 1958-1963. Mitt. Forstl. Bundes-versuchsanst. Mariabrunn, 65:1-48.

Jewett DM; Matsumura F; Coppel HC, 1976. Sex pheromone specificity in the pine sawflies: interchange of acid moieties in an ester. Science, USA, 192(4234):51-53

Johansson BG; Anderbrant O; Simandl J; Avtzis ND; Salvadori C; Hedenstr÷m E; Edlund H; H÷gberg HE, 2001. Release rates for pine sawfly pheromones from two types of dispensers and phenology of Neodiprion sertifer. Journal of Chemical Ecology, 27(4):733-745; 20 ref.

Joly R, 1953. Les lophyres des pins. Revue Forestiere Francaise, 4:269-272.

Jurc M, 2001. Harmful entomofauna (Coleoptera, Lepidoptera, Hymenoptera) on Austrian pine (Pinus nigra Arn.) in Slovenia. In: Proceedings of the 5th Slovenian Conference on Plant Protection, Catez ob Savi, Slovenia, 276-283.

Juutinen P, 1967. Zur Bionomie und zum Vorkommen der Roten Kiefernbuschhornblattwespe (Neodiprion sertifer Geoffr.) in Finnland in den Jahren 1959-65. Communicationes Instituti Forestalis Fenniae, 63.5:1-129.

Juutinen P, 1982. Occurrence and biological control of Neodiprion sertifer in Finland. Allgemeine Forstzeitschrift, No. 8:230-232

Juutinen P; Varama M, 1986. Occurrence of the European pine sawfly (Neodiprion sertifer) in Finland during 1966-83. Folia Forestalia, Institutum Forestale Fenniae, No. 662:39pp.

Kangas E, 1941. Beitrag zur Biologie und Gradation von Diprion sertifer Geoffr. (Hym., Tenthredinidae). Annales Entomologici Fennici, 7:1-31.

Kangas E, 1963. _ber das schSdliche Auftreten der Diprion-Arten (Hym., Diprionidae) in finnischen KiefernbestSnden in diesem Jahrhundert. Zeitschrift fnr angewandte Entomologie, 51:188-194.

Kharitonova NZ; Zhukov VV, 1989. Study of the population characteristics of Neodiprion sertifer to aid the organization of its control. Lesovedenie i Lesnoe Khozyaistvo, No. 24:115-118

Knerer G; Atwood CE, 1973. Diprionid sawflies: polymorphism and speciation. Changes in diapause and choice of food plants led to new evolutionary units. Science, USA, 179(4078):1090-1099

Kolk A; Sierpinski A, 1999. The infestation of Polish forests by insect pests in 1998 and the forecast for 1999. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO Working Party 7.03.10, Sion-Cha^circumflex~teauneuf, Switzerland, 20-23 April, 1999., 89-93.

Kolomiets NG; Stadnitskii GV; Vorontsov AI, 1972. The fox-coloured sawfly (distribution, biology, injuriousness, natural enemies, control measures). Ryzhii sosnovyi pilil'shchik (rasprostranenie, biologiya, vred, estestvennye vragi, mery bor'by). Novosibirsk, Izdatel'stvo Nauka, Sibirskoe Otdelenie. USSR, 148 pp.

Kolonits J, 1965. A Neodiprion sertifer Geoffr. Tletm=dja Ts kßrosftßsa hazßnkban. The bionomics and injuriousness of N. sertifer. ErdTsz. Kutat., 61:225-239.

Krehan H, 1999. Sawfly-gradation in pine stands in Carinthia - different methods of monitoring. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO Working Party 7.03.10, Sion-Cha^circumflex~teauneuf, Switzerland, 20-23 April, 1999., 154-157.

Krieg A, 1955. Untersuchungen nber die Polyedrose von Neodiprion sertifer (Geoffr.). Arch. Ges. Virusforschung, 6:163-174.

Kusevska M, 1973. Dahlbominus fuscipennis Zett. as a regulator of populations of Neodiprion sertifer Geoffr. Godisnik, Sumarki Institut Skopje, 9:65-73.

Kuusik A; Kopvillem H, 1970. Experimental data on the cold resistance of the eggs of Neodiprion sertifer in Estonia. Eesti NSV Tead. Akad. Toimet. (Biol.), 19:329-335.

Larsson S; Bjorkman C; Kidd NAC, 1993. Outbreaks in diprionid sawflies: why some species and not others? In: Wagner MR, Raffa KF, eds. Sawfly life history adaptations to woody plants. San Diego, USA; Academic Press, Inc., 453-483

Leontovyc R; Turcßni M; Z·brik M, 1999. The pests of pine stands in Slovakia. In: Forster B, Knizek M, Grodzki W, eds. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO WP 7.03.10, April 20-23, 1999, Sion-Chateauneuf, Switzerland, 84-88.

Liston AD, 1995. Compendium of European sawflies. Gottfrieding, Germany: Chalastos Forestry.

Lyons LA, 1964. The European pine sawfly, Neodiprion sertifer (Geoff.)(Hymenoptera: Diprionidae). A review with emphasis on studies in Ontario. Proceedings of the Entomological Society of Ontario, 94:5-37.

Lyons LA, 1977. Parasitism of Neodiprion sertifer (Hymenoptera: Diprionidae) by Exenterus spp. (Hymenoptera: Ichneumonidae) in Ontario, 1962-1972, with notes on the parasites. Canadian Entomologist, 109(4):555-564

Lyons LA; Griffiths KJ, 1962. Observations on the development of Neodiprion sertifer (Geoff.) within the cocoon (Hymenoptera: Diprionidae). Canadian Entomologist, 94:994-1001.

LyytikSinen-Saarenmaa P; Anderbrant O; L÷fqvist J; Hedenstr÷m E; H÷gberg HE, 1999. Monitoring European pine sawfly population densities with pheromone traps in young pine plantations. Forest Ecology and Management, 124(2/3):113-121; 32 ref.

LyytikSinen-Saarenmaa P; Tomppo E, 2002. Impact of sawfly defoliation on growth of Scots pine Pinus sylvestris (Pinaceae) and associated economic losses. Bulletin of Entomological Research, 92(2):137-140; 15 ref.

LyytikSinen-Saarenmaa P; Varama M; Anderbrant O; Kukkola M; Hedenstr÷m E; H÷gberg H-E, 2001. Predicting pine sawfly population densities and subsequent defoliation with pheromone traps. In: Liebhold AM, McManus ML, Otvos IS, Fosbroke SLC, eds. Proc. Integr. Manag. Dynam. of For. Def. Insects, Victoria, British Columbia, Canada, Aug. 15-19, 1999. USDA Forest Service, Northeast Research Station, General Technical Report NE-277:108-116.

Magasi LP, 1981. Forest pest conditions in the Maritimes in 1980. Information Report, Canadian Forestry Service, No. M-X-118:ii + 35 pp.

Malinowski H, 1995. Action of insecticides on forest pest insects. V. Effectiveness of insecticides against forest sawfly (Diprionidae) larvae. Prace Instytutu Badawczego Les^acute~nictwa, No. 801-808:23-34; [With English tables. ^italic~Prace^roman~ No. 803]; 14 ref.

Malinowski H, 2002. Activity of azadirachtin against Diprionidae larvae. Sylwan, 146(4):17-24; 19 ref.

Martineau R; LavallTe A, 1975. QuTbec region. Canadian Forestry Service, Annual Report, Forest Insect and Disease Survey, 1974, 37-56.

Martinek V, 1972. Die _bervermehrung der Roten Kiefernbuschhornblattwespe (Neodiprion sertifer (Geoff.)) und die Bedeutung ihrer Parasiten in KnieholzbestSnden. Rozpravy Ceskosl. Akad. Ved, Rada Matematickych a Prirodnich Ved, 82:1-115.

Martinek V, 1974. Cocoon parasites of the European pine sawfly Neodiprion sertifer (Geoff.) in the dwarf-growth stands of Pinus mugo Turra in Bohemia. Rozpravy Ceskosl. Akad. Ved, Rada Matematickych a Prirodnich Ved, 84:1-126.

Martinek V; Kudler J, 1964. Chemical protection of the mountain pine in the Krusne hory Mountains against the feeding of Neodiprion sertifer (Geoffr.) and influence on the parasites. Prace vyzk. ustav. lesn., 29:227-260.

Martini A; Baldassari N; Baronio P, 1999. Gynandromorphism and its manifestations in Diprionid Hymenoptera. Bollettino dell'Istituto di Entomologia `Guido Grandi' della Universita degli Studi di Bologna, No. 53:87-107; 41 ref.

Martini A; Baldassari N; Baronio P; Anderbrant O; Hedenstr÷m E; H÷gberg HE; Rocchetta G, 2002. Mating disruption of the pine sawfly Neodiprion sertifer (Hymenoptera: Diprionidae) in isolated pine stands. Agricultural and Forest Entomology, 4(3):195-201; 21 ref.

McGugan BM; Coppel HC, 1962. Biological control of forest insects, 1910-1958. In: A review of the biological control attempts against insects and weeds in Canada, Pt II. Techn. Commun. No. 2, Commonwealth Institute of Biological Control, Trinidad, 35-216.

McMillin JD; Wagner MR, 1993. Influence of stand characteristics and site quality on sawfly population dynamics. In: Wagner MR, Raffa KF, eds. Sawfly life history adaptations to woody plants. San Diego, USA; Academic Press, Inc., 333-361

Meshkova VL, 1998. Use of insect pathogens in the integrated forest pest management in Ukraine. Methodology of forest insect and disease survey in Central Europe. Proceedings, First Workshop of the IUFRO WP 7.03.10, Ustron^acute~-Jaszowiec, Poland, 21-24 April, 1998., 93-100; 13 ref.

Meshkova VL, 1999. Forest pests outbreaks prognosis on the base of climatic factors analysis [in the Ukraine]. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO Working Party 7.03.10, Sion-Cha^circumflex~teauneuf, Switzerland, 20-23 April, 1999., 74-79; 5 ref.

Mihalache G; Mihalciuc V; Simionescu A; Ianculescu M, 1998. Aspects regarding the phytosanitary state of Romanian forests in the period 1997-98. Methodology of forest insect and disease survey in Central Europe. Proceedings, First Workshop of the IUFRO WP 7.03.10, Ustron^acute~-Jaszowiec, Poland, 21-24 April, 1998., 141-151.

Mihkelson S, 1977. Vastsediapausist punakal mSnnivaablasel. Summary: On the larval diapause of the European pine sawfly. Metsanduslikud Uurimused, 13:234-241.

Mihkelson S, 1980. The role of the egg parasite Achrysocharella ruforum in controlling numbers of the European pine sawfly. Metsanduslikud Uurimused, Estonian SSR, 16:18-25

Monro HAU; Kirby CS, 1963. Fumigation of nursery stock as a possible means of retarding the spread of the European pine sawfly. Canadian Department, Forest Entomology & Pathology Br., Bi-monthly Progtamme Report, 19:2.

Morimoto N; Nakamura G, 1989. Studies on the alpine form of the European pine sawfly, Neodiprion sertifer Geoffroy (Hym.: Diprionidae) in the Central Japanese Alps. I. The life cycle and the mortality process of egg and larval stages. Applied Entomology and Zoology, 24(4):358-371

Morris ON; Cunningham JC; Finney-Crawley JR; Jaques RP; Kinoshita G, 1986. Microbial insecticides in Canada: their registration and use in agriculture, forestry and public and animal health. Bulletin of the Entomological Society of Canada, 18(2 Supplement):43pp.

Nakamura H, 1982. Ecological studies on the European pine sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera: Diprionidae). III. Spatial distribution pattern of larval colonies. Japanese Journal of Applied Entomology and Zoology, 26(1):23-28

NiemelS P; Mannila R; MSntsSlS P, 1982. Deterrent in Scots pine, Pinus sylvestris, influencing feeding behaviour of the larvae of Neodiprion sertifer (Hymenoptera, Diprionidae). Annales Entomologici Fennici, 48:57-59.

Niklas OF; Franz J, 1957. Begrenzungsfaktoren einer Gradation der Roten Kiefernbuschhornblattwespe (Neodiprion sertifer (Geoffr.)) in Sndwestdeutschland 1953 bis 1956. Mitt. Biol. Bundesanst. Land- u. Forstw., Berlin-Dahlem, 89:1-39.

Nuorteva M, 1972. TumamonisSrmi÷viruksen kSyt÷stS ruskean mSntypistiSisen (Neodiprion sertifer Geoffr.) torjunnassa. (Use of the nuclear polyhedrosis virus in the control of the European pine sawfly (Neodiprion sertifer Geoffr.)). Silva Fennica, 6:172-186.

Oehlke J, 1965. Die in europSischen Kiefernbuschhornblattwespen (Diprionidae) parasitierenden Ichneumonidae. Beitrage zur Entomologie, 15:791-879.

Okutani T; Ito T, 1957. On a pine sawfly in high mountains of Japan. The New Entomologist, 6:1-3.

Olofsson E, 1987. Mortality factors in a population of Neodiprion sertifer (Hymenoptera: Diprionidae). Oikos, 48(1):297-303

Olofsson E, 1988. Epizootiology of the nuclear polyhedrosis virus of the European pine sawfly, Neodiprion sertifer. PhD Dissertation, Swedish University of Agricultural Sciences, Uppsala.

Olofsson E, 1988. Persistence and dispersal of the nuclear polyhedrosis virus of Neodiprion sertifer (Geoffroy) (Hymenoptera; Diprionidae) in a virus-free lodgepole pine plantation in Sweden. Canadian Entomologist, 120(10):887-892

Olofsson E, 1989. Oviposition behaviour and host selection in Neodiprion sertifer (Geoffr.) (Hym., Diprionidae). Journal of Applied Entomology, 107(4):357-364

Olofsson E, 1992. Predation by Formica polyctena Forster (Hym., Formicidae) on newly emerged larvae of Neodiprion sertifer (Geoffroy) (Hym., Diprionidae). Journal of Applied Entomology, 114(3):315-319

Opalicki K, 1980. Effects of biological and chemical insecticides on the change in haemocytes in larvae of gypsy moth (Lymantria dispar) and pine sawflies (Diprion pini and Neodiprion sertifer). Glasnik za Sumske Pokuse, No. 20:299-362; 89 ref.

Podgwaite JD; Rush P; Hall D; Walton GS, 1984. Efficacy of the Neodiprion sertifer (Hymenoptera: Diprionidae) nucleopolyhedrosis virus (baculovirus) product, Neochek-S. Journal of Economic Entomology, 77(2):525-528

Pollini A, 1979. Diprionidae injurious to pines. Informatore Fitopatologico, 29(9):19-21

Prop N, 1960. Protection against birds and parasites in some species of tenthredinid larvae. Arch. Neerl. de Zool., 13:380-447.

Pschorn-Walcher H, 1965. The ecology of Neodiprion sertifer (Geoffr.)(Hym.: Diprionidae) and a review of its parasite complex in Europe. Technical Bulletin, Commonwealth Institute of Biological Control, 5:33-97.

Pschorn-Walcher H, 1967. Biology of the ichneumonid parasites of Neodiprion sertifer (Geoffroy)(Hym.: Diprionidae) in Europe. Technical Bulletin, Commonwealth Institute of Biological Control, 8:7-52.

Pschorn-Walcher H, 1970. Studies on the biology and ecology of the alpine form of Neodiprion sertifer (Geoff.) (Hym.: Diprionidae) in the Swiss Alps. Zeitschrift fur Angewandte Entomologie, 66(1):64-83

Pschorn-Walcher H, 1973. Die Parasiten der gesellig lebenden Kiefern-Buschhornblattwespen (Familie Diprionidae) als Beispiel fnr Koexistenz und Konkurrenz in multiplen Parasit-Wirt-Komplexen. Verhandlungen der Deutschen Zoologischen Gesellschaft, 66. Jahresversammlung, Gustav Fischer Verlag, 136-145.

Pschorn-Walcher H, 1982. Suborder Symphyta, sawflies. In: Schwenke W, ed. Die Forstschaedlinge Europas. 4. Hautflügler und Zweiflügler. Hamburg, Germany: Paul Parey, 66-129.

Pschorn-Walcher H, 1988. Die Parasitenkomplexe europäischer Diproinidae in öklogisch-evolutionsbiologischer Sicht. Z. zool. Syst. Evolut.-forsch., 26:89-103.

Pschorn-Walcher H; Eichhorn O, 1973. Studies on the biology and ecology of the egg parasites (Hym.: Chalcidoidea) of the pine sawfly Neodiprion sertifer (Geoff.) (Hym.: Diprionidae) in Central Europe. Zeitschrift fur Angewandte Entomologie, 74:286-318.

Raizenne H, 1957. Forest sawflies of southern Ontario and their parasites. Canadian Department of Agriculture, Publication 1009.

Rieske LK; Townsend L; Anderbrant O; Hedenström E; Högberg HE, 2001. Captures of male European pine sawflies (Hymenoptera: Diprionidae) in pheromone-baited traps in Kentucky. Journal of Entomological Science, 36(1):67-73.

Rivers CF; Crooke M, 1962. Virus control of the sawfly (Neodiprion sertifer Geoff.). Proceedings 5th World Forestry Congress, Seattle (1960), 2:951-952.

Rohwer SA, 1918. New sawflies of the subfamily, Diprioninae (Hym.). Proceedings of the Entomology Society of Washington, 20:79-90.

Ross HH, 1951. Suborder Symphyta (=Chalastogastra). In: Muesebeck CFW, Krombein KV, Townes HK, eds. Hymenoptera of America north of Mexico, synoptic catalog. USDA, Monogr. no. 2:4-88.

Ross HH, 1955. The taxonomy and evolution of the sawfly genus Neodiprion. Forest Science, 1:196-209.

Ryvkin BW, 1957. Die Kiefernblattwespen Wei¯ru¯lands und ihre Parasiten. BeitrSge zur Entomologie, 7:457-482.

Safronov AN, 1996. Protection of Scots pine stands against defoliators. Lesnoe Khozyaistvo, No. 5:49-50.

Sch÷nwiese F, 1935. Beobachtungen und Versuche anlSsslich einer _bervermehrung von Lophyrus sertifer Geoffr. (rufus Panz.) in SndkSrnten in den Jahren 1931/32. Zeitschrift fur Angewandte Entomologie, 21:463-500.

Schaffner JV Jr, 1939. Neodiprion sertifer (Geoff.), a pine sawfly accidentally introduced into New Jersey from Europe. Journal of Economic Entomology, 32:887-888.

Schedl W, 1994. First field experiments with pheromone sticky traps for sawfly control in Austria (Hymenoptera: Symphyta: Diprionidae). Entomologia Generalis, 18(3/4):235-240

Scheidter F, 1934. Forstentomologische BeitrSge. 28. Bestimmungstabelle fnr die Raupen der Gattung Lophyrus (Diprion). Zeitschrift fur Pflanzenkrankheiten und Pflanzenschutz, 44:406-411.

Schwenke W, 1979. On the role of the moulting inhibitor Dimilin in forest protection and forest ecosystems. Anzeiger fur Schadlingskunde Pflanzenschutz Umweltschutz, 52(7):97-102

Schwerdtfeger F, 1936. Zur Kenntnis der roten Kiefernbuschhornblattwespe Diprion sertifer Geoff. (Lophyrus rufus Panz.). Zeitschrift fur Pflanzenkrankheiten und Pflanzenschutz, 46:513-534.

Sezen K; Yaman M; Demirbag Z, 2001. Insecticidal potential of Serratia marcescens Bn10. Biologia (Bratislava), 56(3):333-336; 24 ref.

Simandl J, 1989. Seasonal changes in the synusia of pine sawflies (Hym., Diprionidae) during their latency. Journal of Applied Entomology, 108(3):217-226

Simandl J, 1993. Flight dynamics of males in a non-outbreaking population of the European pine sawfly Neodiprion sertifer (Geoff.) (Hymenoptera, Diprionidae). Lesnictvi^acute~, 39(6):241-243; [With English table and figure]; 13 ref.

Simandl J; Anderbrant O, 1995. Spatial distribution of flying Neodiprion sertifer (Hymenoptera, Diprionidae) males in a mature Pinus sylvestris stand as determined by pheromone trap catch. Scandinavian Journal of Forest Research, 10(1):51-55

Sitowski L, 1925. Do biologii pasorzytow borecznika (Lophyrus Latr.). Roczn. Nauk Rolnicz. i Lesnych, 14:1-25.

Skatulla U, 1975. _ber die Wirkung des Entwicklunghemmers Dimilin auf Forstinsekten. Anzeiger fur SchSdlingskunde, Pflanzenschutz, Umweltschutz, 48:145-147.

Smith DR, 1979. Suborder Symphyta. In: Krombein KV, Hurd PD, Smith DR, Burks BD, eds. Catalog of Hymenoptera in America North of Mexico, 1. Washington DC, USA: Smithsonian Institution Press.

Smith DR, 1993. Systematics, life history, and distribution of sawflies. In: Wagner MR, Raffa KF, eds. Sawfly life history adaptations to woody plants. San Diego, USA; Academic Press, Inc., 3-32

Smits A; Johansson E; Cugala D, 1996. Population dynamics of European pine sawfly Neodiprion sertifer (Fourcroy) (Hym., Diprionidae). Baltic Forestry, 2(1):32-37; 49 ref.

Sullivan CR, 1965. Laboratory and field investigations on the ability of eggs of the European pine sawfly, Neodiprion sertifer (Geoffroy) to withstand low winter temperatures. Canadian Entomologist, 97:978-993.

Suzuki A, 1964. On Neodiprion sertifera [sertifer] Geoffroy. Transl. For. Res. Br. Can. No. 92, 13 pp. Transl. from Japanese Journal of Applied Zoology (Tokyo), 6(5):254-273.

Takeuchi K, 1940. A systematic study on the suborder Symphyta (Hymenoptera) of the Japanese Empire (III). Tenthredo, 3:187-199.

TerSs I, 1982. The climbing activity of larvae of the European pine sawfly, Neodiprion sertifer (Hymenoptera, Diprionidae), knocked from small pines. Notulae Entomologicae, 62:9-12.

Tiihonen P, 1970. Ruskean mSntypistiSisen (N. sertifer Geoffr.) tuhojen vaikutuksesta mSnnik÷iden kasvuun EtelS-Pohjanmaalla, Pohjois-Satakunnassa ja LSnsi-Uudellamaalla vuosina 1960-1967. [_ber die Einwirkungen des Schadfrasses der Roten Kiefernbuschhornblattwespe auf den Zuwachs der KiefernbestSnde im sndlichen Pohjanmaa, im n÷rdlichen Satakunta und im westlichen Uusimaa in den Jahren 1960-67.] Comm. Inst. For. Fenn., 71.3:1-21.

Trewhella KE; Leather SR; Day KR, 1997. The effect of constitutive resistance in lodgepole pine (Pinus contorta) and Scots pine (P. sylvestris) on oviposition by three pine feeding herbivores. Bulletin of Entomological Research, 87(1):81-88.

Trewhella KE; Leather SR; Day KR, 2000. Variation in the suitability of Pinus contorta (lodgepole pine) to feeding by three pine defoliators, Panolis flammea, Neodiprion sertifer and Zeiraphera diniana. Journal of Applied Entomology, 124(1):11-17.

Tsankov G; Rosnev B; Mirchev P, 1980. Results of using a nuclear polyhedral virus preparation for microbiological control of the pine sawfly Neodiprion sertifer in Bulgaria. Gorskostopanska Nauka, 17:75-79.

USDA, 1980. European pine sawfly (Neodiprion sertifer) - Delaware - new state record. Cooperative Plant Pest Report, 5(2):50.

Van Frankenhuyzen F, 2002. Neodiprion sertifer (Geoffroy), European pine sawfly, and N. lecontei (Fitch), redheaded pien sawfly (Hymenoptera: Diprionidae). In: Mason PG, Huber JT, eds. Biological control programmes in Canada, 1981-2000. Wallingford, UK: CAB International, 199-201.

Viitasaari M, 2002. A review of the extant families of the Hymenoptera, Symphyta. In: Viitasaari M, ed. Sawflies (Hymenoptera, Symphyta), I. A review of the suborder, the Western Palearctic taxa of Xyeloidea and Pamphilioidea. Helsinki, Finland: Tremex Press Ltd., 175-195.

Viitasaari M; Varama M, 1987. SahapistiSiset 4. HavupistiSiset (Diprionidae). Summary: Sawflies 4. Conifer sawflies (Diprionidae). Univ. Helsinki, Dep. Agric. and For. Zool., Reports 10:1-79.

Voolma K, 2000. Forest and insects - from pest control to the maintenance of species richness. Akadeemilise Metsaseltsi Toimetised, 11:155-177.

Wagner MR; Raffa KF; eds, 1993. Sawfly life history adaptations to woody plants. San Diego, USA: Academic Press, Inc., 581 pp.

Wallace DR; Sullivan CR, 1973. First successful total laboratory development of eggs of the European pine sawfly (Neodiprion sertifer (Geoff.)). Bi-monthly Research Notes, 29(1):3

Wallner WE, 1968. European pine sawfly control with aircraft application of concentrate insecticidal sprays. J. Econ. Ent., 61:1666-1667.

Watson WY, 1955. Two gynandromorphic sawflies. Bi-monthly Programme Report for Division of Forest Biology, Department of Agriculture, Canada, 11:1.

Wilson LF, 1966. Effects of different population levels of the European pine sawfly on young Scotch pine trees. Journal of Economic Entomology, 59:1043-1049.

Wilson LF, 1971. European pine sawfly. Forest Pest Leaflet, USDA For. Serv. No. 98.

Wright JW; Wilson LF; Randall WK, 1967. Differences among Scotch pine varieties in susceptibility to European pine sawfly. Forest Science, 13:175-181.

Xiao G; Huang X; Zhou S, 1985. The Chinese sawflies of the family Diprionidae (Hymenoptera, Symphyta). Scientia Silvae Sinicae, 21:30-43.

Xiao G; Zhou S; Huang X, 1984. Seven new species of conifer sawflies from China (Hymenoptera: Diprionidae). Entomotaxonomia, 6:141-150.

Yaman M; Nalcacioglu R; Demirbag Z, 2001. Viral control of the European pine sawfly, Neodiprion sertifer (Geoffroy) in Turkey. Turkish Journal of Biology, 25(4):419-425.

Ziogas A; Zolubas P, 1998. Forest pest survey and management in Lithuania. Methodology of forest insect and disease survey in Central Europe. Proceedings, First Workshop of the IUFRO WP 7.03.10, Ustron^acute~-Jaszowiec, Poland, 21-24 April, 1998., 63-67.

Zivojinovic D, 1967. [A contribution to the study of diapause in the pine sawfly (Neodiprion sertifer Geoffr.).] Zastita Bilja, 18(96-97):349-356.

Zivojinovic D; Sidor C, 1975. European pine sawfly (Neodiprion sertifer Geoffr.) and biological control using virus in SR Serbia (Yugoslavia). VIII International Plant Protection Congress, Moscow 1975. Reports and informations. Section V. Biological and genetic control. Moscow. USSR, 224-225

Zolubas P, 1999. Pest status and recent insect outbreaks in pine forests of Lithuania. Methodology of forest insect and disease survey in Central Europe. Proceedings of the Second Workshop of the IUFRO Working Party 7.03.10, Sion-Cha^circumflex~teauneuf, Switzerland, 20-23 April, 1999., 80-83; 5 ref.

Links to Websites

Top of page
WebsiteURLComment
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

Distribution Maps

Top of page
You can pan and zoom the map
Save map