Pineus pini
(pine woolly aphid)

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Identity

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Preferred Scientific Name

• Pineus pini sensu lato

Preferred Common Name

• pine woolly aphid

Other Scientific Names

• Anisophleba pini Koch, 1857
• Aphis pini Gmelin, 1790
• Kermaphis pini var. laevis Maskell, 1885
• Kermes pini Macquart, 1819
• Pineus boerneri Annand, 1928
• Pineus havrylenkoi Blanchard, 1944
• Pineus laevis (Maskell, 1885) Börner, 1907
• Pineus pini (Macquart, 1819) Börner, 1907
• Pineus simmondsi Yaseen & Ghani, 1971
• Pineus sylvestris Annand, 1928

International Common Names

• English: pine, adelgid; pine, aphid, common; pine, aphid, eurasian; pine, woolly, aphid; scots pine, adelges

Local Common Names

• Australia: pine adelgid
• Denmark: fyrrestammelus
• Finland: mäntykirva
• Germany: Wollaus, Europaeische Kiefern-; Wollaus, Kieferntrieb-
• New Zealand: pine adelgid; pine twig chermes
• Norway: furubarlus
• Sweden: tall-lus

EPPO code

• PINEPI (Pineus pini)

Summary of Invasiveness

Top of page P. pini (sensu lato) can cause problems on exotic species of two-needled pines (subgenus Haploxylon) even within its native range, as reported for P. pini from Germany by Steffan (1970b), from France by Gaumont (1978) and from Italy by Covassi and Binazzi (1981).

Infestation by P. pini (sensu lato) causes premature needle shedding and reduction in the length of infested needles, reducing productivity and leading to up to 50% loss of growth increment (Day et al., 2003) and up to 20% tree mortality (Odera, 1974). Stress can make pine trees more vulnerable to attack (Madoffe and Austara, 1993; Day et al., 2003). Damage to trees is most severe under warm, dry conditions (Mailu et al., 1978a).

Wind can facilitate the spread of Pineus (McClure, 1989a), but transmission between countries and between continents is mainly by human transport of infested planting material of two-needled pines (Barnes et al., 1976). The cryptic habit of these insects make them very difficult to detect at plant quarantine inspection, and after introduction they may be overlooked for years because the attention of forest entomologists is often focused on more easily detected pests. In Zimbabwe, the pest was accidentally introduced in 1962 but was not noticed until 1968 (Barnes et al., 1976).

Once introduced and established, alien P. pini (sensu lato) can spread at an average rate of spread of 15 km per year (McClure, 1989b). In Zimbabwe it spread rapidly and reached all the important areas of afforestation within 12 years of its introduction; nearly all the hard pines were attacked with varying degrees of severity, whereas introduced species of soft pine were not (Barnes et al., 1976).

P. pini (sensu lato) has been recorded outcompeting native insects on pines in Kyrgyzstan (Ponomareva and Gabrid, 1981) and in north-eastern North America (McClure 1984a, 1989b).

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Hemiptera
•                         Suborder: Sternorrhyncha
•                             Unknown: Aphidoidea
•                                 Family: Adelgidae
•                                     Genus: Pineus
•                                         Species: Pineus pini

Notes on Taxonomy and Nomenclature

Top of page Damaging populations of asexually-reproducing adelgid aphids belonging to the genus Pineus have been recorded from many temperate and tropical countries around the world in the literature, some under the name Pineus pini, others as P. laevis or P. boerneri, or as one of the other names given in the list of 'non-preferred scientific names'. It should be noted that, in this datasheet, the names in this list are not necessarily taxonomic synonyms. At present there is no completely reliable method for identifying each of the taxa within this complex, at least some of which are taxonomically distinct from each other. P. boerneri is distinct from P. pini, but these species are difficult to differentiate by morphology alone (Blackman and Eastop, 1994). P. havrylenkoi and P. sylvestris are thought to be synonyms of P. boerneri; P. simmondsi Yaseen & Ghani, 1971, recorded from Pinus roxburghii in Pakistan, may also be (Blackman and Eastop, 1994). P. pini (Gmelin) and P. pini (Linnaeus) are synonyms of P. pini (Macquart); Gmelin's names are not regarded as taxonomically valid because he often used trinomials. P. laevis is currently regarded as a synonym of P. pini (Macquart) (Blackman and Eastop, 1994).

Pineus cladogenous Fang & Sun (1985) and Pineus harukawai Inouye (1953), known from Japan, are not regarded as part of the Pineus pini sensu lato complex (Blackman and Eastop, 1994).

Description

Top of page The following is based on Carter (1971). Pineus are such small insects that they are normally prepared as slide mounts so that morphological details can be studied at high magnification. The dorsum bears a regular arrangement of segmental chitinous plates ornamented with wax glands, whose relative sizes and arrangement are of taxonomic importance. In life, the reddish-brown body is usually concealed within a tangle of white wax threads.

Wingless (apterous) adult females are each 0.9-2.3 mm long, the body being ovoid to turbinate; the heavily sclerotized sistens is easier to study than the lightly sclerotized summer form (progrediens); Carter (1971) provides an illustration of the dorsum. Adults are similar to the immature instars, being ovoid with dorsal segmentally arranged sclerotized plates ornamented with wax glands, and short legs and antennae concealed beneath the body. However, the adult females each possess an ovipositor, which is lacking in the immature stages, and the antennae are proportionally very short, with only 1-3 segments that are often partly or completely fused together. There is greater fusion of dorsal plates in adults than in immature Pineus, especially on the head and anterior parts of the thorax.

Winged adult female Pineus resemble other winged aphids in having the body clearly divided into head, thorax and abdomen, and in possessing well-developed legs and antennae; Carter (1971) provides an illustration of the dorsum. The body length is 1.0-1.9 mm. They differ from alate Aphididae in each possessing an ovipositor like that of the apterous female. There are five antennal segments, with each of segments III, IV and V often bearing a single, large sensorium. Compound eyes are present. The shape and venation of the wings is sometimes taxonomically useful.

P. pini (sensu lato) eggs are laid in a nest of white wax fibres secreted by the wax pores of the adult female. Each egg is fairly elongate-ovoid, about 0.2 mm long, and yellow when laid but gradually changes colour to brownish-red as the embryo develops.

First-instar crawlers of Pineus species are very small (each 0.2-0.4 mm long), reddish-brown, ovoid and wingless, with three pairs of legs and small, three-segmented setae; the last antennal segment bears a long apical seta. Eyes are represented only by three oscelli on each side of the head.

Distribution

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P. pini is native to western and Central Europe (Blackman and Eastop, 1994) and P. boerneri may be of eastern Asian origin (Blackman and Eastop, 1994). Records of these species from other parts of the world represent introduced populations.

The geographical records listed for P. pini sensu lato can be tentatively assigned to the following species:

P. pini: Western and Central Europe, Australia, New Zealand.

P. boerneri: China, Taiwan, Hawaii, California, north-eastern USA, Chile and Argentina, eastern and southern Africa, Australia, New Zealand and probably Malaysia; also possibly Pakistan.

Distribution Table

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

History of Introduction and Spread

Top of page Pineus boerneri (as P. pini) was first recorded from Africa (Zimbabwe) in 1962 (Barnes et al., 1976), and from South Africa in 1978 (Zwolinski, 1989). It was probably introduced to Africa accidentally on pine scions from Australia (Barnes et al., 1976).

Risk of Introduction

Top of page P. pini (sensu lato) are very small, cryptic insects that are easily overlooked. They have already been accidentally introduced to a number of countries on infested planting material of two-needled pines, and have subsequently become pests. Cuttings and seedlings of two-needled pines should be subject to very close examination before being exported/imported to a country where P. pini (sensu lato) does not occur.

Habitat

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Pineus boerneri has been recorded preferentially attacking the 3-year-old growth of the lower canopy of Pinus resinosa in the north-eastern USA, feeding beneath bark flakes or at the base of the needle sheaths (McClure, 1982).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
	Terrestrial – Managed	Cultivated / agricultural land	Present, no further details	Harmful (pest or invasive)
		Protected agriculture (e.g. glasshouse production)	Present, no further details	Harmful (pest or invasive)
		Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
		Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
		Disturbed areas	Present, no further details	Harmful (pest or invasive)
		Rail / roadsides	Present, no further details	Harmful (pest or invasive)
		Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
	Terrestrial ‑ Natural / Semi-natural	Natural forests	Present, no further details	Harmful (pest or invasive)
		Natural grasslands	Present, no further details	Harmful (pest or invasive)
		Riverbanks	Present, no further details	Harmful (pest or invasive)
		Wetlands	Present, no further details	Harmful (pest or invasive)
		Cold lands / tundra	Present, no further details	Harmful (pest or invasive)
Littoral
		Coastal areas	Present, no further details	Harmful (pest or invasive)

Hosts/Species Affected

Top of page All the recorded host plants of P. pini sensu lato are members of the subgenus Diploxylon (two-needled pines), not subgenus Haploxylon (pines with needles occurring in groups of more than two) (Anon., 1971). In former Czechoslovakia, with only one exception, infestations of Pineus found on members of Hapoxylon were always Pineus cembrae or P. strobi, not P. pini (Hochmut, 1969).

Pineus boerneri was recorded attacking 39 species of Pinus in Kenya (Odera, 1974), including all the species listed in this datasheet. Host plants that have evolved together with an aphid species have usually evolved defences against aphid infestation that result in slower aphid reproduction on that host.

McClure (1990) suggested that the native host of P. boerneri was Pinus thunbergii; the aphid had significantly greater survival and fecundity on Pinus resinosa (native to America) than on P. thunbergii (native to China).

Host Plants and Other Plants Affected

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Growth Stages

Top of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

Top of page Injury to Pinus resinosa caused by Pineus boerneri in the north-eastern USA included discoloration of foliage, distortion, resinosis, death of branches and death of trees (McClure, 1982). In Africa, other symptoms observed include reduction in the length of infested needles (Day et al., 2003). In Kenya, Mailu et al. (1982b) found that a highly significant negative relationship between needle length and aphid numbers suggested that measurements of needle lengths alone might be adequate to estimate the intensity of infestation by P. boerneri. Damage to trees is most severe under warm, dry conditions (Mailu et al., 1978a).

List of Symptoms/Signs

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Biology and Ecology

Top of page Genetics

Steffan (1970a) discussed the evolution of asexual lines in species of adelgids including P. pini, and remarked that the asexual strain often had a different number of chromosomes to the sexual strain. P. pini may have arisen as an asexual strain of P. orientalis (Marchal, 1911; Carter, 1969, 1971).

Blackman et al. (1995) found that different populations of Pineus ?boerneri varied in their chromosome number, as might be expected of an asexual species. Specimens with 2n=16 were recorded from Hawaii and California, USA; samples with 2n=17 were recorded from California, Africa, Australia and New Zealand. Some samples from Australia and New Zealand had 2n=19 and 2n=21. Blackman et al. (1995) postulated that the 17-chromosome form of P. boerneri could have arisen from the 16-chromosome form through a single dissociation of one of the longest pair of chromosomes. Such dissociations are common in permanently parthenogenetic populations of aphids (Blackman, 1980).

Physiology and Phenology

In North America, Pineus boerneri is parthenogenetic, trivoltine and passes through six stages of development (McClure, 1989b); however, the generation overlap was so pronounced that all the developmental stages were present at almost all times of the year.

The following is based on Carter (1971). The life cycle of P. pini (sensu lato) differs from that of most adelgids by its simplicity. There may be a partial migration of alatae (sexuparae) to Picea (the ancestral primary host) each year, but males are never produced and sexual reproduction and gall formation on Picea does not occur.

In the absence of sexual reproduction, gall-forming females (gallicolae) do not occur, and asexual reproduction by wingless and winged (alate) females continues throughout the year. There are at least three overlapping generations annually. Clutches of eggs are laid (one egg at a time) over a period of several weeks, resulting in considerable differences between the hatching times of the first and last eggs laid. There are five immature instars before an adult is produced. Overwintering is as wingless adult females (hiemosistentes) that are heavily sclerotized, and as second- and third-instar larvae. In spring, the larvae develop into apterous progredientes (the summer form of the adult female, which is lightly sclerotized) and alate (winged) females.

Reproductive Biology

Reproduction is asexual all year, by the parthenogenetic production of eggs.

Environmental Requirements

Stress can make pine trees more vulnerable to attack by P. boerneri (Madoffe and Austara, 1993; Day et al., 2003). Air pollution was found to make pines more susceptible to attack by P. pini in Finland, possibly due to a resultant reduction in predatory mite populations (Heliovaara and Vaisanen, 1989).

In the north-eastern USA, survival of P. boerneri was higher on branches than on needles during winter, but was higher on needles than on branches in spring and summer (McClure, 1984a). Wind facilitated the spread of P. boerneri in north-eastern North America, and extremes of summer and winter temperatures determined the ultimate limits of its latitudinal distribution; heavy rain reduced its abundance (McClure, 1989a).

In Malawi, the biological performance and pest status of P. boerneri is influenced by external factors especially the within-tree and seasonal fluctuations in the level of nitrogen, total rainfall and canopy structure (Chilima and Leather, 2001). Prolonged, heavy rainfall reduced population levels significantly, mainly through eggs and crawlers being washed off the tree, whereas numbers increased in hot, dry weather; other causes of mortality were found to be excessive heat and crawler dispersion (Mailu et al., 1980).

In the UK, Carter and Barson (1973) found that the threshold temperature for flight by winged Pineus was 16°C, and at least 64% of the catch was taken in the afternoon; fluctuations in the mean date of flight each year were probably related to the phonological condition of the food plant.

Associations

In Scotland, UK, P. pini is sometimes associated with a covering of the fungus Cucurbitaria pithyophila [Curreya pityophila] on the lower branches of Pinus sylvestris, occasionally killing the bark (Murray and Parry, 1969).

In north-eastern America, where the introduced P. boerneri and the native P. coloradensis occur together on Pinus resinosa, P. boerneri was found to maintain numerical dominance over P. coloradensis (McClure, 1984a) and displaced P. coloradensis in the majority of stands over a 5-year period (McClure, 1989b). However, where P. boerneri and Matsucoccus resinosae (an introduced scale insect) occurred together, the presence of M. resinosae on the branches resulted in significantly reduced numbers of P. boerneri on both branches and needles, whereas the presence of P. boerneri had no significant effect on numbers of M. resinosae. In one-third of the affected stands, P. boerneri was excluded by the scale insect within 3 years (McClure, 1990).

Natural enemies

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Notes on Natural Enemies

Top of page Introduced populations of P. boerneri have been successfully controlled by the introduction of predatory species of Leucopis. The most effective control agent recorded is L. tapiae in Hawaii, where Leucopis tapiae (as L. obscura, on Maui, Molokai and Kauai islands) and L. nigraluna (on Hawaii island) were found to maintain P. boerneri (as P. pini) populations below economically significant levels (Nakao et al., 1978; Culliney et al., 1988).

In Kenya, predatory Exochomus spp. were found to remove 12% of the Pineus boerneri population (Mailu et al., 1980). In Zimbabwe, coccinellid beetles were responsible for a marked decline in the infestation (Anon., 1971).

Biological control attempts have also been made in Australia (Waterhouse and Sands, 2001). Only Exochomus quadripustulatus, imported from the UK, became established there. The native predator Diomus pumilo is considered an important predator.

Means of Movement and Dispersal

Top of page Natural Dispersal

Dispersal of P. pini (sensu lato) is mainly achieved by the tiny first-instar larvae, which walk actively and seek new feeding sites. They are capable of walking a few metres only, but can be picked up and carried over longer distances by the wind. Wind facilitated the spread of P. boerneri in north-eastern North America (McClure, 1989a). Transmission between countries and between continents is by human transport of infested planting material (Barnes et al., 1976).

Crawlers of P. boerneri were found to be positively phototactic in some circumstances, but were negatively phototactic and positively thigmotactic when seeking to settle and feed, therefore colonizing tight crevices; branch undersides were settled much more often than better-lit branch upper surfaces (Mailu et al., 1982a).

Occasionally, in crowded conditions, P. pini (sensu lato) produce winged females that fly to seek new host trees. In the UK, winged Pineus were recorded mainly in late May and June each year; the threshold temperature for flight by winged Pineus was 16°C, and at least 64% of the catch was taken in the afternoon; fluctuations in the mean date of flight each year was probably related to the phenological condition of the food plant (Carter and Barson, 1973).

Pathway Vectors

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Plant Trade

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Impact Summary

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Economic Impact

Top of page Pineus boerneri

In sub-Saharan Africa pine trees are widely grown, both commercially and by small-scale farmers; they are an important source of building material and fuelwood. Infestation by Pineus boerneri causes premature needle shedding and reduction in the length of infested needles, reducing productivity and leading to up to 50% loss of growth increment (Day et al., 2003) and up to 20% tree mortality (Odera, 1974). In South Africa, Zwolinski (1990) recorded loss of increment at breast height, and overall height, in Pinus pinaster trees at 63 and 65%, respectively. Madoffe and Austara (1990) recorded P. boerneri (as P. pini) causing over 20% loss of dry matter in seedlings of Pinus patula in Tanzania. Similarly, infestation of developing cones of P. pinaster in South Africa reduced overall seed yield by 23%, not counting those cones that failed to reach maturity (Zwolinski et al., 1989). Murphy (1996) estimated that P. boerneri and another introduced pine aphid (Eulachnus rileyi) together were causing a growth increment loss of US $1.5 million annually. Damage to trees is most severe under warm, dry conditions (Mailu et al., 1978a). Survey procedures and criteria for assessment of damage by P. boerneri were given by Mailu et al. (1980).

P. boerneri was recorded injuring and killing trees at all of the infested sites studied in Connecticut and Massachusetts (north-eastern USA) (McClure, 1989b). It is listed (as P. havrylenkoi) as one of the main pests of pine trees in Argentina (Fiorentino and Diodato de Medina, 1991).

Mendel et al. (1994) recorded P. pini from Jordan for the first time, causing severe damage to young stands of Aleppo and stone pines. It is not clear whether this record referred to P. boerneri or P. pini.

In Australia (Victoria), Minko (1962) mentioned P. pini causing damage to Pinus radiata in nurseries; this may actually have referred to P. boerneri.

Pineus pini (sensu lato)

In response to problems with adelgids (including P. pini) on exotic conifers grown in the numerous tree nurseries east of Oldenburg, Germany, Steffan (1970b) recommended that these tree species should not be planted in shelter belts, parks or forests within the growing region.

In Sweden, infestation by P. pini of pine tree hosts in glasshouse cultures of chanterelle mushrooms limited long-term cultivation of this crop (Zambonelli, 2002).

Covassi and Binazzi (1981) mentioned that, in Italy, P. pini is injurious to young pine trees in nurseries, new plantations, parks and gardens in several parts of Italy.

In the Republic of Georgia, a study of damage caused by P. pini (sensu lato) found that infestation could result in a 32% reduction in annual height increment, and various other signs of damage, and concluded that the adelgid was a serious threat to pine trees in that zone (Saradzhishvili, 1985).

In Kyrgyzstan, P. pini (sensu lato) (as P. pini) outcompeted the native Oligonychus piceae in the same ecological niche; local natural enemies were ineffective against it, suggesting it was an introduction, and Pineus numbers were only reduced after Leucopis spp. were released (Ponomareva and Gabrid, 1981).

Canakcioglu (1972) listed P. pini as one of the most important forest pests in Turkey.

Diagnosis

Top of page Pineus are such small insects that they are normally prepared as slide mounts so that morphological detail can be studied at high magnification. Carter (1971) provides a method for the preparation of slide mounts and keys to identify the genus Pineus and to more-or-less separate the four species that occur in the UK; the key covers P. pini and P. orientalis, but not P. boerneri.

Detection and Inspection

Top of page Strong light and magnification are required, and the bark and needle sheaths need to be destructively examined. P. pini (sensu lato) occur as brownish-red ovoids in nests of white wax fibres beneath bark flakes and bud scales, and concealed within the needle sheaths. Clusters of yellow or brownish-red eggs may also be present in the wax nests; the eggs are not attached to the host plant.

Similarities to Other Species/Conditions

Top of page P. pini is morphologically indistinguishable from P. orientalis, a species that reproduces sexually and alternates between Picea (the primary host) and Pinus sylvestris. At present, P. boerneri is not reliably morphologically distinguishable from P. pini and P. orientalis because morphological variation in these species has not been sufficiently well analysed and no identification key is available.

Prevention and Control

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Phytosanitary Measures

Restricted movement and thorough inspection of planting material of Haploxylon pines is the most effective way to prevent P. pini (sensu lato) entering a country.

Cultural Control and Sanitary Methods

Stress can make pine trees more vulnerable to attack by P. boerneri (Madoffe and Austara, 1993); avoidance of marginal growing conditions through more attention to the selection of suitable planting sites can avoid problems with Pineus infestation (Day et al., 2003).

Possible management of P. boerneri in Malawi, through the regulation of nitrogen nutrition and canopy structure, was discussed by Chilima and Leather (2001).

In response to problems with adelgids (including P. pini) on exotic conifers grown in the numerous tree nurseries east of Oldenburg, Germany, Steffan (1970b) recommended that these tree species should not be planted in shelter belts, parks or forests within the growing region.

Host-Plant Resistance

Different species of pine tree, and different provenances of any one pine species, vary in their susceptibility to attack by P. pini (sensu lato) (Simpson and Ades, 1990b; Falkenhagen, 1991a, b). The level of susceptibility of a particular pine species may vary also between one locality and another, reflecting different levels of environmental stress between sites. Simpson and Ades (1990b) called for more research on the planting of multi-clonal mixtures and the pattern in which they are planted, to minimize losses.

Replanting with species or provenances of pines less susceptible to attack is an option for the control of P. pini (sensu lato), but is a long-term and possibly expensive solution that may incur hidden costs in reduced yield or quality, e.g. Pinus radiata has been replaced with the less susceptible P. patula in East Africa, but the latter species is inferior in fibre quality (Day et al., 2003).

Odera (1974) mentioned that Pinus ayacahuite and P. strobes var. chiapensis are resistant to attack by Pineus boerneri (as Pineus pini) in Kenya.

Biological Control

Where members of P. pini (sensu lato) have been accidentally introduced, they are good candidates for classical biological control (Mills, 1990). Pineus boerneri has been controlled successfully by introduced natural enemies in Hawaii, New Zealand and Chile.

Introduced populations of P. boerneri have been successfully controlled by the introduction of predatory species of Leucopis. The most effective control agent recorded is L. tapiae in Hawaii, where Leucopis tapiae (as L. obscura, on Maui, Molokai and Kauai islands) and L. nigraluna (on Hawaii island) were found to maintain P. boerneri (as P. pini) populations below economically significant levels (Nakao et al., 1978; Culliney et al., 1988).

In 1991 L. tapiae was also found to have accidentally entered Malawi, but has not provided such effective control there as in Hawaii (Greathead, 1995; Day et al., 2003). An account of various (mostly unsuccessful) introductions of alien predator species to East Africa against P. boerneri is given by Day et al. (2003).

In Kyrgyzstan, P. pini (sensu lato) (as P. pini) outcompeted the native Oligonychus piceae in the same ecological niche; local natural enemies were ineffective against it, suggesting it was an introduction, and Pineus numbers were only reduced after Leucopis spp. were released (Ponomareva and Gabrid, 1981).

Introduction of the predatory bug Tetraphleps raoi to Kenya in 1975 resulted in some reduction in P. boerneri populations, but not sufficient to prevent reduction of tree growth and occasional tree mortality (Karanja and Aloo, 1990).

References

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Annand PN, 1928. A contribution toward a monograph of the Adelginae (Phylloxeridae) of North America. Stanford University Publications, University Series, 6(1):1-146.

Anon, 1971. [I] The Pine woolly aphid [Pineus pini]: present position. [II] A further note on the Pine woolly aphid. Research Newsletter, Rhodesia Forestry Commission; 1973, No. 6; 7, 5-6; 1-2.

Anon, 1978. Forest and shade trees - new host records. United States Department of Agriculture, Cooperative Plant Pest Report, 3:39, 561.

Barnes RD, Jarvis RF, Schweppenhauser MA, Mullin LJ, 1976. Introduction, spread and control of the pine woolly aphid, Pineus pini (L.), in Rhodesia. Suid-Afrikaanse Bosboutydskrif, No. 96:1-11

Blackman RL, 1980. Chromosome numbers in the Aphididae and their taxonomic significance. Systematic Entomology, 5(1):7-25

Blackman RL, Eastop VF, 1994. Aphids on the world's trees: an identification and information guide. Wallingford, UK: CAB International.

Blackmann RL, Watson GW, Ready PD, 1995. The identity of the African pine woolly aphid: a multidisciplinary approach. Bulletin OEPP, 25(1/2):337-341; 10 ref.

Blanchard EE, 1944. Descripciones y anataciones de afidoideos argentinos. Acta Zoologica Lilloana, 2:15-22.

Börner C, 1907. Systematik und Biologie der Chermiden. Zoologischer Anzeiger, 32:413-420.

Canakcioglu H, 1972. The occurrence of aphid species (Aphidoidea) in Turkish forests. Anzeiger fur Schadlingskunde und Pflanzenschutz, 45(10):152-154; 22 ref.

Carter CI, 1969. Three species of adelgids (Homoptera, Adelgidae) new to Britain. Ent. mon. Mag. 105 (1262/1264), (167-9). [8 refs.].

Carter CI, 1971. Conifer woolly aphids (Adelgidae) in Britain. Bull. For. Comm., Lond. No. 42 pp. 51 + 21 plates. [50 refs. Price 75p.].

Carter CI, Barson G, 1973. Flight activity of alate Adelgids (Homoptera, Aphidoidea) in southern England. Bulletin of Entomological Research, 62(3):507-516

Chilima CZ, 1991. The status and development of conifer aphid damage in Malawi. Workshop Proceedings. Exotic aphid pests of conifers: a crisis in African forestry, Muguga, Kenya: Kenya Forest Research Institute and Food and Agriculture Organization of the United Nations, 64-67.

Chilima CZ, Leather SR, 2001. Within-tree and seasonal distribution of the pine woolly aphid Pineus boerneri on Pinus kesiya trees. Agricultural and Forest Entomology, 3(2):139-145; 46 ref.

Covassi M, Binazzi A, 1981. Contributions to the knowledge of aphids of conifers. IV. Notes on some species of adelgids found in Italy (Homoptera Adelgidae). Redia, 64:303-330

Culliney TW, Beardsley JW Jr, Drea JJ, 1988. Population regulation of the Eurasian pine adelgid (Homoptera: Adelgidae) in Hawaii. Journal of Economic Entomology, 81(1):142-147

Day RK, Kairo MTK, Abraham YJ, Kfir R, Murphy ST, Mutitu KE, Chilima CZ, 2003. Biological control of homopteran pests of conifers in Africa. Pineus boerneri. In: Neuenschwander P, Borgemeister C, Langewald J, eds. Biological Control in IPM Systems in Africa. Wallingford, UK: CABI Publishing, 104-106.

Falkenhagen ER, 1991. Provenance variation in Pinus muricata in South Africa. Silvae Genetica, 40(2):50-57; 15 ref.

Falkenhagen ER, 1991. Provenance variation in Pinus radiata at six sites in South Africa. Silvae Genetica, 40(2):41-50; 18 ref.

Fang SY, Sun JH, 1985. Study on Korean pine adelgids in Northeast China, with descriptions of two new species. Journal of North-East Forestry University, China, 13(3):1-7

Fiorentino DC, Diodato de Medina L, 1991. Brief survey of the forest insect pests of Argentina. Investigacion Agraria, Sistemas y Recursos Forestales Madrid, Spain; Instituto Nacional de Investigacion y Tecnologia Agraria y Alimentaria, No. 16:181-190

Gabrid NV, 1982. Feeding specialisation of the pine adelgid Pineus pini L. (Macq.). Entomologicheskie Issledovanie v Kirgizii, No. 15:62-76

Gaumont R, 1978. Practical tables for determining the principal forms of the Chermesidae (= Adelgidae) of France. Revue Forestiere Francaise, 30(1):21-36

Greathead DJ, 1995. The Leucopis spp. (Diptera: Chamaemyiidae) introduced for biological control of Pineus sp. (Homoptera: Adelgidae) in Hawaii: implications for biological control of Pineus ?boerneri in Africa. Entomologist, 114(2):83-90

Heliovaara K, Vaisanen R, 1989. Invertebrates of young Scots pine stands near the industrialized town of Harjavalta, Finland. Silva Fennica, 23(1):13-19

Hochmut R, 1969. Occurrence of woolly aphids of the genus Pineus on different Pine species in the Bolevec arboretum, Czechoslovakia. Anz. Schadlingsk. 42 (6), (90-2). [4 refs.].

Inouye M, 1953. Monographische Studie ueber die japanischen Koniferen-Gallenlaeuse (Adelgidae). Bulletin of the Sapporo Government Forest Experiment Station, 15:1-91.

Jaskiewicz B, Sawinska A, 2005. Aphids (Homoptera, Aphidodea) inhabiting the shrubs of Pinus mugo Turra in the green areas of Lublin. Part I. The population dynamics. Acta Agrobotanica, 58(1):153-164.

Karanja MK, Aloo TC, 1990. The introduction and establishment of Tetraphleps raoi Ghauri as a control of woolly aphid in Kenya. Technical Note - Kenya Forestry Research Institute, No. 12:i + 11 pp.

Koch CL, 1857. Die pflanzenläuse Aphiden getreu nach dem Leben abgebildet und beschrieben. Nürnburg, 275-336.

Macquart MJ, 1819. Notice sur les insects hémiptères du genre Psylle. Séances Publiques de la Société d’amateurs des Sciences et Arts de la ville de Lille. Séance du Mars, 1819:81-86.

Madoffe SS, Austar+ O, 1993. Abundance of the pine woolly aphid Pineus pini in Pinus patula stands growing on different sites in the Sao Hill district, Tanzania. Commonwealth Forestry Review, 72(2):118-121; 35 ref.

Madoffe SS, Austara , 1990. Impact of pine woolly aphid, Pineus pini (Macquart) (Hom., Adelgidae), on growth of Pinus patula seedlings in Tanzania. Journal of Applied Entomology, 110(4):421-424

Mailu AM, Khamala CPM, Rose DJW, 1978. Evaluation of pine woolly aphid damage to Pinus patula and its effect on yield in Kenya. East African Agricultural and Forestry Journal, 43(3):259-265

Mailu AM, Khamala CPM, Rose DJW, 1980. Population dynamics of pine woolly aphid, Pineus pini (Gmelin) (Hemiptera: Adelgidae), in Kenya. Bulletin of Entomological Research, 70(3):483-490

Mailu AM, Khamala CPM, Rose DJW, 1982. Establishment of pine woolly aphid, Pineus pini (Gmelin) (Adelgidae) on some host trees in Kenya. Kenya Journal of Science and Technology, (Biological Sciences), 3(2):61-68

Mailu AM, Khamala CPM, Rose DJW, 1982. Sampling techniques for populations of pine woolly aphid, Pineus pini (Gmelin) (Adelgidae). Kenya Journal of Science and Technology, (Biological Sciences), 3(1):9-18

Mailu AM, Rose DJW, Khamala CPM, 1978. Sequential surveys for the pine woolly aphid, Pineus pini (L.) Homoptera: Adelgidae in Kenya. East African Agricultural and Forestry Journal, 43(3):252-258

Marchall P, 1911. La spanadrie et l’obliteration de la reproduction sexuée chez les Chermes. Compte rendu hebdomadiare des Scéances de l’Academie des Sciences, Paris, 153:299-302.

Maskell WM, 1885. Note on an aphidian insect infesting pine trees, with observations on the name "Chermes" or "Kermes". Transactions and Proceedings of the New Zealand Institute (1884), 17:13-29.

McClure MS, 1982. Distribution and damage of two Pineus species (Homoptera: Adelgidae) on red pine in New England. Annals of the Entomological Society of America, 75(2):150-157

McClure MS, 1984. Influence of cohabitation and resinosis on site selection and survival of Pineus boerneri Annand and P. coloradensis (Gillette) (Homoptera: Adelgidae) on red pine. Environmental Entomology, 13(3):657-663

McClure MS, 1984. Pineus boerneri Annand (Homoptera: Adelgidae): a new or another record from the People's Republic of China? Proceedings of the Entomological Society of Washington, 86(2):460-461.

McClure MS, 1989. Biology, population trends, and damage of Pineus boerneri and P. coloradensis (Homoptera: Adelgidae) on red pine. Environmental Entomology, 18(6):1066-1073

McClure MS, 1989. Importance of weather to the distribution and abundance of introduced adelgid and scale insects. Agricultural and Forest Meteorology, 47(2-4):291-302

McClure MS, 1990. Cohabitation and host species effects of the population growth of Matsucoccus resinosp (Homoptera: Margarodidae) and Pineus boerneri (Homoptera: Adelgidae) on red pine. Environmental Entomology, 19(3):672-676

Mendel Z, Assael F, Saphir N, Zehavi A, Kafisheh W, 1994. New distribution records of Matsucoccus josephi and Pineus pini (Homoptera) on pine trees in parts of the Near East. Phytoparasitica, 22(1):9-18

Mills NJ, 1990. Biological control of forest aphid pests in Africa. Bulletin of Entomological Research, 80(1):31-36

Minko G, 1959. An adelgid on Monterey Pine in the Ovens Plantations [, Victoria]. For. tech. Pap. For. Comm. Vict. No. 1, (19-23). 8 refs.

Minko G, 1962. Insects observed attacking Pinus radiata D. Don. in nurseries. For. tech. Pap. For. Comm. Vict. No. 10, (34-47). 2 refs. Limited distribution.

Murphy ST, 1996. Status and impact of invasive conifer aphid pests in Africa. Impact of diseases and insect pests in tropical forests. Proceedings of the IUFRO Symposium, Peechi, India, 23-26 November 1993., 289-297; 16 ref.

Murray JS-and-Parry, WH, 1969. The association of Pineus pini and Cucurbitaria pithyophila on Scots Pine in northern Scotland. Scot. For. 23 (1), (8-13). [3 refs.].

Nakao HK, Funasaki GY, Higa SY, Lai PY, 1978. Introductions for biological control in Hawaii - 1977 and 1978. Proceedings of the Hawaiian Entomological Society, 23(3):425-430

Odera JA, 1974. The incidence and host trees of Pine woolly aphid, Pineus pini (L.), in East Africa. Commonwealth Forestry Review, 53(2):128-136; 10 ref.

Ponomareva RE, Gabrid NV, 1981. Competitive interaction between the pests of common pine. In: Aukshtikal'nene AM, ed. Noveshie dostizheniya lesno entomologii (po materialam USh s"ezda VEO, Vil'nyus, 9-13 oktyabrya 1979 g.) Institut Zoologii i Parazitologii Akademii Nauk Litovsko SSR Vilnius USSR, 136-138

Questienne P, 1979. Notes sur quelques insectes nuisibles aux pins au Maroc. Annales de Gembloux, 85(2):113-130.

Ravensberg K, 1954. Jaarb. Proefsta. Boomkwek. Boskoop 1953:17-21 (in Dutch).

Saradzhishvili KG, 1985. The damage caused by Pineus pini in the subalpine zone of Soviet Georgia. Lesnoe Khozyaistvo, No. 12:46-48

Shakachite O, Chenduauka B, 1991. A report on current status of pine insect pests with particular reference to pine aphids in Zambia. In: Ceisla WM, Odera J, Cock MJW, Ceisla PM, eds. Workshop Proceedings: exotic aphid pests of conifers. A crisis in African forestry, Muguga, Kenya 3-6 June 1991. Rome, Italy, Food and Agriculture Organization (FAO), 85-93.

Simpson JA, Ades PK, 1990. Screening Pinus radiata families and clones for disease and insect pest resistance. Australian Forestry, 53(3):194-199

Simpson JA, Ades PK, 1990. Variation in susceptibility of Pinus muricata and Pinus radiata to two species of Aphidoidae. Silvp Genetica, 39(5-6):202-206

Steffan AW, 1970. The introduction and spread of Adelgidae attacking conifers in the tree-nursery region of the Ammerland. Z. Angew. Ent. 65 (3), (345-56). [23 refs.].

Steffan AW, 1970. The origin of anholocyclic, parthenogenetic species of Adelgidae on the basis of a study of life-cycle forms and their cytology. Z.angew.Ent. 65 (4), (444-52). [38 refs.].

Toccafondi P, Covassi M, Pennacchio F, 1991. Studies on the entomophagous predators of scale insects of the genus Matsucoccus Cock. in Italy. II Bio-ethological notes on Rhyzobius chrysomeloides (Herbst) in pine forests of Liguria (Coleoptera: Coccinellidae). Redia, 74(2):599-620

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

Wojtan R, Bystrowski C, Tomusiak R, Zasada M, 2006. Assessment of the health condition of pine stands following the pine woolly aphid (Pineus pini L.) outbreak in the uków and Ostrów Mazowiecka Forest Districts. (Ocena stanu zdrowotnego drzewostanów sosnowych po gradacji ochojnika sosnowego (Pineus pini L.) w nadlesnictwach uków i Ostrów Mazowiecka.) Sylwan, 150(8):11-19. http://sylwan.ibles.waw.pl/

Yaseen M, Ghani MA, 1971. Descriptions, and notes on the biology, of four new species of Adelgidae from West Pakistan. Bull. ent. Res. 61 (2), (191-205). [7 ref.].

Yuronis VA, 1984. Thirty-one species of aphids (Aphidinea) new to the Lithuanian SSR, found in 1981-1982. In: Ionaitis, ed. Novye i redkie dlya Litovskoi SSR vidy nasekomykh. Soobshcheniya i opisaniya 1984 goda. Vilnius, USSR; Institut Zoologii i Parazitologii Akademii Nauk Litovskoi SSR, 13-17.

Zambonelli A, 2002. Current research on chanterelle cultivation in Sweden. In: Danell E, Hall I, eds. Edible mycorrhizal mushrooms and their cultivation. Proceedings of the Second International Conference on Edible Mycorrhizal Mushrooms, Christchurch, New Zealand, 3-6 July 2001, 0-4.

Zondag R, 1977. Pine twig chermes or pine woolly aphid. Forest and Timber insects in New Zealand No. 25.

Zwolinski JB, 1989. The pine woolly aphid, Pineus pini (L.) a pest of pine in South Africa. South African Forestry Journal, No. 151:52-57

Zwolinski JB, 1990. Preliminary evaluation of the impact of the pine woolly aphid on condition and growth of pines in the southern Cape. South African Forestry Journal, No. 153:22-26

Zwolinski JB, Grey DC, Mather JA, 1989. Impact of pine woolly aphid, Pineus pini (Homoptera: Adelgidae) on cone development and seed production of Pinus pinaster in the Southern Cape. South African Forestry Journal, No. 148:1-6

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