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379© Springer International Publishing Switzerland 2016
T.D. Paine, F. Lieutier (eds.), Insects and Diseases of Mediterranean Forest
Systems, DOI 10.1007/978-3-319-24744-1_13
Chapter 13
Invasive Insects in the Mediterranean
Forests of Chile
Sergio A. Estay
Abstract The Mediterranean ecosystem of Chile comprises the most populated
region of the country as well as the most important passageways for international
trade. This has caused the rate of introductions of exotic insects to be the highest of
the country. As a consequence, it has been observed an increasing number of exotic
insects currently being established in three different situations: natural forests, com-
mercial forest plantations and urban trees.
In natural forests, the most serious threat is the invasion of Cinara cupressi , caus-
ing severe damage on the endangered native Cupressaceae, Austrocedrus chilensis.
Apart from this case, records of exotic insects attacking native trees are scarce. On
the other hand, forest plantations of exotic species, mainly Pinus radiata and
Eucalyptus spp ., have been affected for several insect invasions in the last years.
These new invasions represent a challenge for the Chilean forest industry, making
necessary the adoption of new silvicultural techniques and sanitary protocols for
exporting wood products. Besides commercial plantations, insect invasions are also
taking place in urban areas: poplars, elms and other ornamental trees have under-
gone the introduction of several exotic insects. The effect of this pests has been
perceived as a major threat for the environmental quality of cities, and public opin-
ion is increasingly demanding actions from the government to minimize their
effects.
Considering the increasing rate of detection of exotic insects in Chile, it becomes
clear that invasive insects represent a current threat for Chilean forests and immedi-
ate actions are needed from the government, private companies and society to mini-
mize their negative consequences.
S. A. Estay (*)
Instituto de Ciencias Ambientales y Evolutivas, Facultad de Ciencias , Universidad Austral de
Chile , Casilla 567 , Valdivia , Chile
Center of Applied Ecology and Sustainability (CAPES), Facultad de Ciencias Biológicas ,
Universidad Católica de Chile , Casilla 114-D , Santiago , Chile
e-mail: sergio.estay@uach.cl
380
13.1 Introduction
13.1.1 Mediterranean Chile
The Mediterranean region of Chile represents one of the world’s intensively
exploited Mediterranean ecosystems. With a rapid economic growth, and a increased
centralization of human activities, this ecosystem has suffered signifi cant changes
in structure and disturbance regimes over the past last decades (Fuentes et al. 1993 ;
Rundel 1998 ; Armesto et al. 2007 ; Schulz et al. 2010 ). In Chile, Mediterranean
forests are located between 30°S and 38°S Lat (Donoso 1993 ). This region com-
prises the three largest metropolitan areas of the country: Santiago city, and the
conurbations Valparaiso-Viña del Mar and Concepción-Talcahuano, summing
nearly 13.5 million people (INE 2014 ).
Topographically, this region contains two parallel mountain ranges oriented from
north to south: the Coastal and the Andes mountain ranges. They are separated by a
narrow basin, which is an 80–100 km wide tectonic depression named Central
Depression. The climate is semiarid with hot and dry summers and cold and wet
winters with most precipitation concentrated between April and September (Luebert
and Pliscoff 2006 ). Considering Chile as a rather narrow country, in this article all
geographic descriptions are given in latitude only.
A high fl oristic richness and diversity of plant communities is present in this
region resulting from the high heterogeneity in topography, moisture, soil types and
nutrient availability (Arroyo et al. 1993 , 1995 ). Several and contrasting natural for-
est types are distributed following the North–South and Coast-Mountain gradients.
In the northern-coastal part of the region we may found the Olivillo Forest. This is
a relict forest dominated by Olivillo, Aextoxicon punctatum , a common species of
the south of Chile, which is immersed in a matrix of xerophytic vegetation (Armesto
et al. 2007 ).
The emblematic genus Nothofagus is also present in Mediterranean Chile. In the
highest peaks of the Coastal mountain range, around the city of Santiago, there are
fragments of the deciduous N. macrocarpa . Besides, it is also possible to fi nd mixed
stands of deciduous Nothofagus species in a formation locally known as the Maulino
forest (San Martín and Donoso 1996 ). At both slopes of the Coastal and Andes
mountain range, evergreen sclerophyllous vegetation dominates the landscape
(Armesto et al.
2007 ). According to Armesto et al. ( 2007 ) this formation, dominated
by evergreen trees and shrubs, is the most common plant formation in central Chile.
Xerophytic open woodlands typically associated with grazing pastures, with the
physiognomic aspect of savannas (Fuentes et al.
1990 ), are widespread (Armesto
et al. 2007 ). Above 1500 masl, an upland sclerophyllous woodland called Andean
Montane Woodlands is present. Finally, above treeline and about 2300 masl, the
Montane Woodland is replaced by a low subalpine shrubland (Arroyo et al. 1981 ).
Scattered patches of the conifer Austrocedrus chilensis (Cupressaceae) also occur
near the treeline at various locations between 32°S and 36°S (Rundel 1981 ). Others
forest types, like Palm and Swamp Forests, are also present in this region, although
S.A. Estay
381
limitedly distributed. For further details the reader can consult to Armesto et al.
( 2007 ) and Luebert and Pliscoff ( 2006 ).
Immersed in this matrix of natural ecosystems and human settlements are the
commercial forest plantations. The most important species of these plantations are
the exotic Pinus radiata and two Eucalyptus species, E. nitens and E. globulus .
From these, Pinus radiata is the most important one and the basis for the forestry
industry of the region. Currently, Chile has 1.5 million ha of P. radiata plantations
established across different site types and climate conditions ranging from 30°S to
43°S Lat. (INFOR 2014 ). Eucalyptus plantations sum 800,000 ha located between
29°S and 45°S Lat. (INFOR 2014 ).
Finally, it is important to highlight the presence of urban environments. Many
exotic tree species can be found all over the country in parks and streets. These trees
are highly appreciated by citizens and have a direct impact on the neighborhoods’
appraisal within the city (Gutiérrez 2011 ).
13.1.2 Propagule Pressure
Currently, habitat fragmentation, changes in the land use and pollution are impor-
tant processes in the Mediterranean Chile. Furthermore, this region constitutes the
most important passageway for international trade in and out of the country.
Therefore, the propagule pressure for the introduction of exotic species is likely to
be high (Estay et al. 2012 ). As a matter of fact, several exotic insect species have
already been reported in the region (Grooves and Di Castri 1991 ; Rundel et al. 1998 ;
Ruz 2002 ; Grez et al. 2010 ). However, the way in which these human-related habi-
tat modifi cations are affecting the diversity of native and exotic insect species, and
whether their establishment can be unequivocally associated to human-related
activities are still unknown (but see Estay et al. 2012 ).
Due to the high concentration of shipping ports, Mediterranean Chile undergoes
a high pressure of exotic insect introductions. Ferrada et al. ( 2007 ) and Ide et al.
( 2014 ) estimated this pressure using the number of intercepted exotic insects in
wood articles in Chilean ports. 1,440 interceptions of live insects were registered
between 1996 and 2009. From these, 17.6 % were classifi ed as quarantine forest
pests and 80 % of the interceptions took place in the Mediterranean region (Ide et al.
2014 ).
This high propagule pressure is mostly represented by insects of the order
Coleoptera, accounting for the 86.5 % of the interceptions (Ide et al. 2014 ). It is
particularly important the high rate of insect interceptions from the Cerambycidae
and Scolytidae families. Both comprise several species known to be harmful for
coniferous trees like Pinus spp . Regarding the origin of the interceptions, Ferrada
et al. ( 2007 ) and Ide et al. ( 2014 ) showed that most of the invasive insects came
from Brazil, India, China, Spain and USA.
13 Invasive Insects in the Mediterranean Forests of Chile
382
13.2 Invasive Insects in the Mediterranean Chile
13.2.1 Natural Forests
Despite the high propagule pressure described for the Mediterranean Chile, there
are few records of exotic insects attacking native trees, and only in few cases the
magnitude of the attack may be considered severe enough to represent a threat to
natural forests. For example, Barriga et al. ( 1993 ), after almost 10 years of sampling
natural and artifi cial landscapes along Chile (between 26°S and 38°S), did not detect
any exotic xylophagus Coleoptera on native plants. A similar situation is described
by Fuentes-Contreras et al. ( 1997 ) in their review of aphids in Chile. These authors
listed the presence of just two exotic species associated to native trees, and in both
cases the records are anecdotal. However, from this short list, two cases deserve
special attention: Cinara cupressi and Nematus oligospilus .
13.2.1.1 Cinara cupressi
The cypress aphid, Cinara cupressi (Hemiptera: Aphididae), is a well known pest of
Cupressaceae, damaging natural forests and plantations worldwide (FAO 2009 ).
Currently, it is accepted that C. cupressi is a complex of several anatomically similar
species (Watson et al. 1999 ). FAO ( 2009 ) indicates that the Cinara species presents
in Chile is C. cupressivora. However , this classifi cation is still controversial, and in
this review the name C. cupressi will be used hereafter.
The origin of the species is unclear. Following Watson et al. ( 1999 ), the species
becoming a pest in several countries corresponded to C. cupressivora, original from
the region between eastern Greece and the Caspian Sea. Nevertheless, the origin of
the remainder of the complex ( C. cupressi sensu lato ) is North America. Despite
this unclear origin, the cypress aphid has caused serious damage to natural and
planted forests in Africa, Europe, Latin America and the Caribbean, and the Near
East (FAO 2009 ).
Adult cypress aphids are typically 2–5 mm length, dark brown and covered with
a powdery wax (FAO 2009 ). Colonies typically have between 20 and 80 nymphs
and winged and non-winged adults, located at the host’s branches (FAO 2009 ).
Adults and nymphs suck the plant sap retarding new growth and causing desiccation
and progressive dieback until death on heavily infested trees (Ciesla 1991 ). A sec-
ondary problem caused by aphid feeding is the copious quantities of honeydew,
which promotes the growth of sooty mold (Ciesla 1991 ). The cypress aphid have
been described attacking several genera in the Cupressaceae family like Cupressus,
Juniperus, Thuja, Chamaecyparis, among others, and some authors suggest that any
Cupressaceae could be a suitable host (FAO 2009 ).
Several authors have pointed out the high economic and environmental impacts
of the cypress aphid. In Africa, economic losses has been quantifi ed in Kenya,
where over 2 years 12 % of the trees were killed (Orondo and Day 1994 ). Also in
S.A. Estay
383
the southern and eastern African region the cypress aphid caused losses for US$27.5
million in 1991 with a loss in annual growth increment of US$9.1 million per year
(Murphy et al. 1996 ). In terms of environmental impacts, a major menace is the
impact on the threatened Widdringtonia species in Africa. According to Chapman
( 1994 ), aphids attack are a key component of the pool of menaces acting over this
trees.
In 2003 is detected for fi rst time in Chile, in the district of Pica (20.5°S, 69.3°W,
González et al. 2010 ; Montalva et al. 2010 ) and, by the end of 2008, its presence
was confi rmed across the whole country infesting native and exotic species of
Cupresssacea (Peña and Altmann 2009 ; González et al. 2010 ; Montalva et al. 2010 ).
Aphid’s most important threath is its adaptation to Austrocedrus chilensis , a conifer
species endemic to Chile and Argentina classifi ed as “vulnerable” (Hechenleitner
et al. 2005 ). In the year 2005, 52 % of the 48 survey stations on A. chilensis forests
detected the presence of C. cupressi , and the prevalence increased to 74 % by 2007
(INFOR 2008 ; Peña and Altmann 2009 ). Field data collected in 2005 and 2008 for
three administrative regions of Mediterranean Chile (between 34°S and 38°S Lat.)
showed that 42 %, 5 %, and 10 %, respectively, of the surveyed A. chilensis trees
exhibited >50 % loss of and necrotic foliage combined (Peña and Altmann 2009 ).
Mortality, depending on the region, was 0.2–5 % of trees by the year 2005 in central
Chile (34.5°S–38°S Lat., INFOR 2008 ; Peña and Altmann 2009 ). In 2007, C.
cupressi was detected attacking isolated trees of another emblematic native conifer,
Fitzroya cupressoides (González et al. 2010 ; Montalva et al. 2010 ).
To minimize the impact of C. cupressi on A. chilensis , several governmental
agencies promote a pest management program with emphasis on biological control.
During 2004, Pauesia juniperorum (Hymenoptera: Braconidade) was found parasit-
izing C. cupressi , suggesting this parasitoid enter the country together with C.
cupressi . Unfortunately, the effi ciency of this parasitoid seems to be low (González
et al. 2010 ; Montalva et al. 2010 ). Subsequent introductions of Pauesia sp . from
Spain and Italy have been reported unsuccessful (González et al. 2010 ).
13.2.1.2 Nematus oligospilus
The willow sawfl y (Hymenoptera: Tenthredinidae) is a wasp native of the Holartic
Region that feed on willows (Salix), and since 1980 has been introduced in many
localities in the Southern Hemisphere (Koch and Smith 2000 ). It is a green wasp of
5–7 mm (female) or 4.7–5.3 mm (male) length (Koch and Smith 2000 ), and strictly
parthenogenetic in the Southern Hemisphere (Gonzalez 1989; Caron et al. 2014 ).
This species was fi rst detected in central Chile in 1984, feeding on the exotic
weeping willow ( Salix babylonica ) and the native pencil willow ( S. humboldtiana )
(Gonzalez et al. 1986 ). Currently the willow sawfl y is distributed from 28°S to 40°S
in Chile, covering a large part of the distribution of S. humboldtiana (Gonzalez
1989; Klein–Koch and Waterhouse 2000 ; Estay 2004 ). Attacks on S. humboldtiana
are specially severe in central Chile, causing complete defoliation of trees (Gonzalez
1989; Klein–Koch and Waterhouse
2000 ).
13 Invasive Insects in the Mediterranean Forests of Chile
384
Control strategies are not clearly established, because chemical, cultural and bio-
logical control seems to be ineffi cient and in some cases showing strong collateral
effects (Alderete et al. 2010 ). In particular, parasitoids observed in South America
have shown a very low effi cacy (Alderete et al. 2010 ).
13.2.1.3 Other Exotic Insects of Native Forests
There are few other reports of exotic insects attacking native trees in Mediterranean
Chile and none in the magnitude of the two previous cases. The fi rst one in this list
is the olive scale, Saissetia oleae (Hemiptera: Coccidae), introduced to Chile in the
nineteenth century and nowadays widespread in Mediterranean Chile. This scale
has been reported feeding on several native trees such as Maytenus boaria,
Kageneckia oblonga, Luma apiculata, or Raphithamnus spinosus , but in any case
the damages can be considered a real threat for natural populations of these trees
(Gonzalez 1989 ; González and Lamborot 1989 ; Artigas 1994 ). Aspidiotus nerii is
another scale reported attacking native trees. According to Artigas ( 1994 ), this spe-
cies has been found on several native species like Prosopis chilensis, Beilschimiedia
miersii, Peumus boldus and Cryptocaria alba . Despite the economic importance of
this scale in fruit cultivation, A. nerii is considered just locally important for some
stands of P. chilensis (Klein–Koch and Waterhouse 2000 ). In the order Coleoptera,
the Powder- post beetle, Lyctus brunneus , a cosmopolitan pest of dead-wood, has
been occasionally reported feeding in the wood of some native species, but they
could be easily misidentifi ed with L. chilensis (Rojas and Gallardo 2004 ).
Some other anecdotal records can also be found in literature. Two Hemiptera
have been reported on native trees: Phylloxera similans on Nothofagus alpina
(Carrillo and Cerda 1987 ; Klein–Koch and Waterhouse 2000 ), and Aphis nasturtii
on Drymis winteri (Eastop et al. 1997 ; Fuentes–Contreras et al. 1997 ). Both records
correspond to particular observations and no information about damage is available.
Klein–Koch and Waterhouse ( 2000 ) indicate the presence of the Megachile rotun-
data (Hymenoptera) cutting leaves of Persea lingue , but at a negligible rate.
Lanfranco ( 2010a ) describe the presence of Orgya antiqua (Lepidoptera:
Lymantriidae) on native plants, but without reporting the host species.
A special note deserves the yellowjacket wasp, Vespula germanica. Although
this is not an insect exclusively associated to forests, its impact on native insects,
agriculture, apiculture, wildlife, and recreation activities in national parks is sub-
stantial (Beggs 2001 ; Curkovic et al. 2004 ; Kasper 2004 ). In Chile, this insect was
introduced more than 30 years ago (Edwards 1976 ; Archer 1998 ; Estay and Lima
2010 ), showing at the present time high densities, especially in Central Chile, where
it has become a real threat for recreational and touristic activities, causing also seri-
ous damage to fruit farms and vineyards (Curkovic et al. 2004 ; Estay and Lima
2010 ).
S.A. Estay
385
13.2.2 Forest Plantations
Chilean forestry industry is based on exotic species. Currently Chile has 2.4 million
of ha of planted forest, 88 % immersed in the Mediterranean ecosystem (INFOR
2014 ). 61 %, 22 % and 10 % correspond to plantations of Pinus radiata, Eucalyptus
globulus and Eucalyptus nitens , respectively. In this section we will review the
major insect pests of these three species.
13.2.2.1 Invasive Insects of Pinus radiata
As the same time that P. radiata plantations turned into the main resource of the
Chilean forestry industry, at the beginning of the twentieth century, concerns about
its phytosanitary conditions started to arise. However, it is only at the end of that
century that occurred the major and, still today, more mediatic introduction of an
exotic forest insect. Rhyacionia buoliana (Lepidoptera: Tortricidae) was detected in
Southern Chile in 1980, and today is distributed between 32°S and 42°S Lat
(Lanfranco 2010). Native of Europe, this moth destroy pine shoots causing deformi-
ties such as forked or crooked stems, bushy growth, multiple tops and a lowered
timber quality (Harris and Wood 1967 ; Miller et al. 1961 ). The severity of the dam-
age seems to increase from north to south of Chile (Klein–Koch and Waterhouse
2000 ). Alzamora et al. ( 2002 ) quantifi ed the economic losses in Southern Chile
between 2 % and 30 % of the expected land value depending on the quality of the
forest site. To minimize the impact of this pest, several actions were taken during the
1980s in an unprecedented effort of the government, companies and research cen-
ters. This effort capitalized into the development of new management schemes,
monitoring programs and several control programs. Chilean government, through
the Agricultural and Livestock Service (SAG) and the National Forest Service
(CONAF), concentrated control efforts in the importation of natural enemies, start-
ing a biological control program using Orgilus obscurator (Hymenoptera:
Braconidae). On this regard, Lanfranco (2010) pointed out that followed a success-
ful establishment an initial control, the effi ciency of O. obscurator as biological
control has decreased in the last decade, reaching in some localities, values not
higher than 40 %.
Taking a jump in time to the present day, we found the introduction to Chile of
Sirex noctilio (Hymenoptera: Siricidae). Probably this is the major threat to P. radi-
ata plantations in the last 20 years. The Sirex woodwasp is native of Europe, Asia,
and North Africa, where it is a minor pest (Ciesla 2003 ). However, this species
turned into a serious pest of pines in New Zealand, Australia, South Africa,
Argentina, Uruguay, Brazil and Chile, where it has become one of the most eco-
nomically signifi cant pests of softwood forestry (Villacide and Corley 2012 ).
Damage occurs when females oviposit eggs into stressed or suppressed trees, along
with a phytotoxic mucus and a wood decay fungus ( Amylostereum areolatum )
(Madden 1988 ; Carnegie et al. 2006 ). Trees are drilled by larvae and soon die due
13 Invasive Insects in the Mediterranean Forests of Chile
386
to the combination of the mucus and fungus (Ciesla 2003 ; Carnegie et al. 2006 ). In
Chile, this insect was fi rst detected in January 2001 in the Valparaiso Region (32.5°S
Lat.), and almost simultaneously, a second detection occurred in Los Lagos Region
(40.4°S Lat.), being both events different events of introduction (Rojas and Beèche
2010 ; Beèche et al. 2012 ). Today this pest is under offi cial control by the Agricultural
and Livestock Service, however, its current distribution cover most of the commer-
cial plantations of P. radiata (SAG 2015 ). Suppression strategies currently applied
in Chile consist in cultural and biological control, using the nematode Deladenus
siricidicola and Megarhyssa nortoni (Hymenoptera: Ichneumonidae), combined
with restriction to the movement of wood from infested areas (Rojas and Beèche
2010 ; Beèche et al. 2012 ).
Another important group of exotic pests in Pinus plantations in Chile are the
bark beetles (Coleoptera: Scolytidae). Three species belong to this group: Hylurgus
ligniperda, Hylastes ater and Orthotomicus erosus , all of them detected in Chile
during the 1980s (Ciesla 1988 ; Lanfranco et al. 2002 ). The three species are native
of Eurasia and Africa. Hylastes ater was fi rst detected in Chile in 1983, H. ligni-
perda in 1985 and O. erosus in 1986. Hylurgus ligniperda and H. ater are now dis-
tributed from 33°S to 41°S Lat., and Orthotomicus erosus shows a more restricted
distribution in Chile, between 35°S and 41°S Lat. (Lanfranco et al. 2002 ). The three
species invade fresh stumps and use this material as breeding sites (Ciesla 1988 ;
Lanfranco et al. 2002 ). Adults feed on the root collars and in roots of 1 and 2-year-
old pines, and in case of heavy attacks, they can cause mortality of seedling (Ciesla
1988 ; Lanfranco et al. 2002 ).
Many other exotic insects have established in pine plantations in Chile, but with
minor or no economic impact. Among Coleoptera species, Buprestis novenmaculata
was detected in Chile in 1940, and nowadays is distributed between 32°S and 40°S
Lat. This insect is native from Europe and Northern Africa, but its impact on com-
mercial plantations is negligible because it is associated with deadwood (Lanfranco
et al. 2002 ). On suppressed or dead trees it is also possible to fi nd Ernobius mollis
(Anobiidae) and Xyleborinus saxeseni (Scolytidae). Both species were introduced
to Chile in the 1950–1960s, and currently they are distributed between 32°S and
37°S (Gonzalez 1989; Lanfranco et al. 2002 ). In the order Hemiptera, two species
deserve mention: Pineus borneri (adelgidae) and Eulachnus rileyi (Aphididae). The
North American P. borneri is a long resident in Chile. Some authors suggest that
their introduction would have occurred at the beginning of the twentieth century
(Olalquiaga 1952 ). At the present time, this species is widespread in Chile, although
it seems to be more abundant above 40°S Lat. (Klein–Koch and Waterhouse 2000 ).
The second Hemiptera species, the European E. rileyi was fi rst reported in Chile in
1990 (Cerda and Beeche 1989 ), but probably its introduction occurred long before.
No clear description of its current distribution is available, but putting several refer-
ences and personal communications together, it can be inferred its current distribu-
tion range from 35°S to 42°S Lat. (Fuentes–Contreras et al. 1997 ; Klein–Koch and
Waterhouse 2000 ). Finally, another Siricidae, Urocerus gigas from Eurasia, was
discovered in Chile in the 1970s (FAO
2008 ). Currently, it is distributed between
32°S and 42°S Lat. without causing major damage (Lanfranco et al. 2002 ).
S.A. Estay
387
13.2.2.2 Invasive Insects of Eucalyptus Plantations
If we analyze the history of forest pest introductions in the last 20 years, we can
notice that more introductions of Eucalypt pests than pine pests have occurred in the
Mediterranean Chile. The major problems are caused for two species of the
Australian genus Phoracantha (Coleoptera: Cerambycidae): P. semipunctata and P.
recurva . The former was offi cially detected in Chile in 1973, however, there is an
individual collected in Chile in 1931, which is deposited in the collection of the
Museum of the University of Concepción (Artigas 1994 ). Phoracantha recurva , on
the other hand, was fi rst detected in Chile in 1997 (Lanfranco and Dungey 2001 ;
Beéche et al. 2003 ). Both species are serious pests of Eucalyptus plantations located
in arid environments or in poorly irrigated sites (Artigas 1994 ; Lanfranco and
Dungey 2001 ). Klein–Koch and Waterhouse ( 2000 ) report that the current distribu-
tion of P. semipunctata is between 18°S and 40°S Lat while P. recurva is distributed
from 32°S to 34.5°S Lat. Forest industry and the government has focused manage-
ment efforts in two directions. First, towards an improvement of silvicultural tech-
niques, specially linked to irrigation, in order to avoid stressed individuals
susceptible to attacks (Lanfranco and Dungey 2001 ). Second, towards biological
control using Avetianella longoi (Hymenoptera: Encyrtidae), which is now success-
fully established in Chile achieving variable levels of parasitism depending on loca-
tion (Beéche et al. 2003 ).
Immediately after the detection of P. recurva , Gonipterus scutellatus was discov-
ered on Eucalyptus trees in central Chile (Beéche et al. 1999 ; Estay et al. 2002 ).
Since their introduction, this defoliator has been recognized as a major threat for E.
globulus plantations, specially due to its great adaptation to the climatic conditions
of the Mediterranean Chile, where is able to produce three to four generations per
year (Estay et al. 2002 ). According to Klein-Koch and Waterhouse ( 2000 ), G. scu-
tellatus is the most serious Eucalyptus pest in central Chile. Today the species is
widespread from 32°S to 40°S Lat. (Elgueta 2010 ). Governmental agencies have
introduced Anaphes nitens and Anaphes tasmaniae (Hymenoptera: Mymaridae) as
biological control with signifi cant success (Beéche et al. 1999 ). A recent study sug-
gest that G. scutellatus is actually a complex and that the species already present in
New Zealand, Spain, California and Chile is G. platensis , native of Tasmania
(Mapondera et al. 2012 ). Nevertheless, more studies are necessaries to clarify the
taxonomy of this group.
In the order Hymenoptera, two introduced insects deserve special mention. The
fi rst one is Ophelimus eucalypti (Eulophidae). Native of Australia, this gall-forming
insect was introduced in 2003 and now is widespread in Mediterranean Chile
(Carnieletto 2006 ; SAG 2006 ). However, its impact on Eucalyptus plantations does
not seem to be of relevance. The second one is Leptocybe invasa (Eulophidae),
recently introduced (2014) and restricted to central Chile (32–33°S Lat.). Native of
Australia, it is considered a serious pest of young trees and saplings (SAG 2014 ).
No information is available of its current situation in Chile. Finally, two other
Hemiptera have been reported to Chile, but with very different implications.
Blastopsylla occidentalis (Psyllidae) was reported in 2000 in central Chile (32°S,
13 Invasive Insects in the Mediterranean Forests of Chile
388
Burckahardt and Elgueta 2000 ), but it does not represent a sanitary problem for
Eucalyptus plantations. On the other hand, another Psyllidae, Ctenarytaina euc-
alipty represents a serious pest of E. globulus sapling and young trees. It was intro-
duced in Chile in 1999 (Olivares 2000 ; Goycoolea et al. 2002 ), and currently is
widespread in Chile at least from 20°S to 40°S Lat (Rodrígues and Sáiz 2014 ). Due
to the importance of this pest, governmental and private agencies developed a bio-
logical control program using Psyllaephagus pillosus (Hemiptera: Encyrtidae)
reported as successful (Goycoolea et al. 2002 ).
13.2.3 Invasive Insects of Urban Trees
Urban environments in Chile concentrate the major diversity of exotic insects.
Gardens, parks and other human-made plantations form a mosaic of refugees and
alternative food sources for insects and other small sized organisms, enhancing the
richness of exotic species. Using a extensive sampling for 2 years in the Metropolitan
Region of Chile (33°S), Estay et al. ( 2012 ) reported 81 exotic insect species on
trees, more than 30 % of the identifi ed species.
The diversity of urban trees in Chilean Mediterranean cities has facilitated the
establishment of many well-known insect pests in recent times. Considering the
high number of reported exotic pests in urban environments, this review will focus
on the most important and notable examples.
Glycaspis brimblecombei (Hemiptera: Psylidae) is an Australian insect pest of
several Eucalyptus species such as E. camaldulensis, E. tereticornis and E. rudis
(Valente and Hodkinson 2009 ). This insect was introduced to Chile in 2002, and at
present is distributed from 30°S to 38°S Lat. (Ide et al. 2006 ). In Chile its main host
has been E. camaldulensis , a species commonly used in urban parks. Its damage is
impressive since trees get covered by the cone-shaped protection of nymphs (lerp).
Successive attacks caused defoliation and weakening facilitating the attack of sec-
ondary pests like Phoracantha spp . (Estay 2004 ; Ide et al. 2006 ). Immediately after
introduction, governmental agencies planned the introduction of natural enemies,
which was fi nally carried out in 2003 with the importation of Psyllaephagus luteus
(Hymenoptera: Encyrtidae). However, this effort had variable success depending on
location (Ide et al. 2006 ).
Another defoliator causing serious damage in elms is Xanthogalerucella luteola
(Coleoptera: Chrysomelidae). Although it was reported in Chile as early as 1982
(Askevold 1991 ), it was not until 1994 when this pest was formally reported as
established in Chile (SAG 2005 ). This insect is easily observable in elms in all
Chilean Mediterranean cities between 32°S and 38°S Lat. causing serious defolia-
tions. Considering the level of damage, authorities have started the implementation
of chemical control (Estay 2004 ; SAG 2010 ). This insect is able to develop four
S.A. Estay
389
generations per year in central Chile, amplifying the severity of its damage (Huerta
et al. 2011 ).
Poplars are widely used in urban streets and also as sherterbelts in rural and
semirural landscapes. These trees have undergone the attack of several insects, but
probably the more important ones are Tremex fuscicornis (Hymenoptera: Siricidae)
and Chaitophorus leucomelas (Hemiptera: Aphididae). Tremex fuscicornis is a
wood borer wasp native of Asia and Eastern Europe, where is not considered a pest
(Parra 2010 ). In Chile was detected in 2000 on Populus nigra , but probably its intro-
duction occurred at least 2 years earlier (Parra 2010 ). Nowadays, its impact on
urban trees is high and its damage is observable on several hosts like poplars, wil-
low, black locust, ashleaf maple or common walnut (Estay 2004 ; Parra 2010 ). Its
current distribution span from 32°S to 37°S Lat. with one generation per year (Parra
2010 ). A natural enemy, Megarhyssa praecellens (Hymenoptera: Ichneumonidae),
was introduced simultaneously with the pest, but its effectiveness has been reported
only about 30 % mainly due to its low dispersal ability (Parra 2010 ). To compensate
for this, continuous releases of this parasitoid are needed. Cultural control strategies
are effective to prevent the attack of T. fuscicornis such as irrigation or destruction
of infested trees by burning or chipping (Estay 2004 ; Parra 2010 ). The second
important pest for poplars, C. leucomelas, was fi rst described as present in Chile in
1995 (Muñoz and Beéche 1995 ). Aphid nymphs feed on leaves and buds of poplar
causing in some cases earlier leaves fall (Estay 2004 ). However, the main problem
associated to this species in Central Chile is the abundant production of honeydew
that stains sidewalks, houses, cars, benches or any other object located below the
tree (Estay 2004 ) causing annoyance on citizens. It is currently distributed from 18°
to 36° (Klein–Koch and Waterhouse 2000 ). Although there was an intent of biologi-
cal control in 2001 using Adialytus salicaphis (Hymenoptera: Braconidae), this was
reported as failed (Stanković et al. 2015 ).
Native to Perú and Bolivia Schinus molle is a very common urban tree in
Santiago, capital of Chile. It has been suggested that this tree was introduced to
Chile at the time of the Inca Empire (Silva et al. 2005 ). At present almost all
trees are infested with Calophya schini (Hemiptera: Calophyidae). This small
insect causes small galls in the leaves, sometimes dozen per leafl et (Downer
et al. 1988 ; Estay 2004 ; Zina et al. 2012 ). This massive presence of galls com-
pletely deform leaves and provoke a grayish appearance on infested trees, fol-
lowed by extensive foliage drop (Downer et al. 1988 ; Zina et al. 2012 ). Despite
the high visibility of the damage, there is no information of its ecology on
Mediterranean Chile.
As previously mentioned, there are tens of other exotic insects on urban trees in
Mediterranean Chile, but it is out of the scope of this review to check all of them.
For some of them, simply there is no literature available. On spite of this, Table 13.1
offer to the reader information about some other species present in urban trees of
Chile. For a more complete overview, readers may consult Estay ( 2004 ).
13 Invasive Insects in the Mediterranean Forests of Chile
390
Table 13.1 Exotic insects present in the urban environments of Mediterranean Chile not discussed
in the main text
Order Family Species Hosts
References
for Chile
Coleoptera Buprestidae Melonophila picta Populus sp. Estay
(
2004 )
Coleoptera Scolytidae Scolytus rugulosus Ulmus sp.,
Crataegus sp.
Betula sp.
Elgueta and
Marvaldi
(
2006 )
Hemiptera Aleyrodidae Siphoninus phillyreae Catalpa
bignonioides,
Cercis
siliquastrum,
Magnolia
grandifl ora,
Fraxinus
excelsior,
Ligustrum
vulgare, Punica
granatum, among
others
Muñoz and
Beéche
(
1995 )
Hemiptera Aphididae Drepanoshipum
oregonensis
Acer sp. SAG ( 2010 )
Hemiptera Aphididae Essigella californica Pinus sp.,
Pseudotsuga
menziesii
SAG ( 2004 )
Hemiptera Aphididae Myzocallis boerneri Quercus suber,
Q. ilex
SAG ( 2005 )
Hemiptera Aphididae Tinocallis saltans Ulmus sp.
Hemiptera Coccidae Parthenolecanium
corni
Robinia
pseudoacacia,
Fraxinus
excelsior, Ulmus
sp., Acer sp.,
Crataegus sp.,
Platanus sp.,
Salix babylonica,
Tilia sp., among
others
Gonzalez
(1989)
Hemiptera Diaspididae Carulaspis minima Several
Cupressaceae
Estay
(
2004 )
Hemiptera Diaspididae Hemiberlesia rapax Ficus sp.,
Gleditsia
triacanthos,
Melia azedarach,
Robinia
pseudoacacia,
Acer sp., Schinus
molle,
Pyttosporum sp.,
among others
Gonzalez
(1989)
(continued)
S.A. Estay
391
13.3 Conclusions
Our review allows us to make some generalizations about the status of invasive
insects in Chile. It seems that native forests have not yet experienced a high rate of
introductions of exotic insects. However, the few successful introductions have been
highly harmful. Cinara cupressi represents a major threat to native endangered
conifers, and recognizing that eradication is probably not a realistic option, the con-
servation of these conifers will require a major effort that considers continuing
monitoring and the development of a new biological control program using more
effective natural enemies.
Forest plantations have been affected for the introduction of exotic insects.
Assuming the increasing exchange of goods, the most likely future scenario shows
new introductions of forest pests as harmful as the currently established in Chile.
This scenario (will) require that the forest industry develop new policies such as a
better coordination with governmental agencies, improvements in contingency
plans and an increased investment in the necessary science and technology required
to minimize the impact of these current and future pests.
Society’s demands for a healthy environment incorporate more and more the
sanitary condition of urban vegetation. Citizen’s concern about the impact of forest
Table 13.1 (continued)
Order Family Species Hosts
References
for Chile
Hemiptera Diaspididae Lepidosaphes beckii Magnolia
grandifl ora, Acer
sp., Eleagnus sp.
Gonzalez
(1989)
Hemiptera Diaspididae Parlatoria pittospori Pittosporum sp.,
Pinus sp.,
Phoenix sp.,
Acacia sp.,
Chamaecyparis
lawsoniana,
among others
Estay
(
2004 )
Hemiptera Pseudococcidae Pseudococcus
calceolariae
Schinus molle,
Pinus sp.,
Abutilon sp.,
Ficus sp., Salix
sp, among others
Gonzalez
(1989)
Hemiptera Psyllidae Psyllopsis fraxinicola Fraxinus
excelsior and F.
monophylla
Burckhard
(
1994 )
Hemiptera Thaumastocoridae Thaumastocoris
peregrinus
Eucalyptus sp. Ide et al.
(
2011 )
Hemiptera Tingidae Corythuca ciliata Platanus sp,
Fraxinus sp.,
Tilia sp.
Estay
(
2004 )
13 Invasive Insects in the Mediterranean Forests of Chile
392
pests on urban trees is a social phenomenon that arises in the last decade in Chile.
These demands are an opportunity for joining efforts with citizens, local groups or
municipalities into invasive species programs. “Citizen Science” has demonstrated
to be very useful for conservation science, and could be a signifi cant step forward
for monitoring and early alert systems.
This review will be incomplete without mention the effects of global change.
Several studies suggest an important reduction of precipitations in Mediterranean
Chile, which will affect tree’s health. This scenario will promote outbreaks and
range expansion of current forest pests, but will also facilitate the establishment of
opportunistic insects. The development of national policies to ameliorate these
impacts is an urgent task.
References
Alderete M, Liljesthrom G, Fidalgo P, Alderete M, Liljesthrom G, Fidalgo P (2010) Bio–ecología
y perspectivas para el manejo de la avispa sierra del sauce, Nematus oligospilus , Manejo
Integrado de Plagas Forestales n° 10. INTA, Bariloche
Alzamora R, Apiolaza LA, Ide S (2002) Evaluación física y económica de pérdidas en volumen
debido al daño de Rhyacionia buoliana (Schiff.) en plantaciones de Pinus radiata (D. Don) en
la Novena y Décima Regiones de Chile. Bosque 23:29–42
Archer M (1998) The world distribution of the Euro–Asian species of Paravespula (Hym, Vespine).
Entomol Mon Mag 134:279–284
Armesto JJ, Arroyo MTK, Hinojosa LF (2007) The Mediterranean environment of Central Chile.
In: Veblen TT, Young KR, Orme AR (eds) The physical geography of South America. Oxford
University Press, New York, pp 184–199
Arroyo MTK, Armesto JJ, Villagrán C (1981) Plant phenological patterns in the high Andean
Cordillera of central Chile. J Ecol 69:205–223
Arroyo MTK, Armesto JJ, Squeo F et al (1993) Global change: the fl ora and vegetation of Chile.
In: Mooney H, Fuentes E, Kronberg B (eds) Earth–system responses to global change: con-
trasts between North and South America. Academic Press, New York, pp 239–264
Arroyo MTK, Cavieres L, Marticorena C et al (1995) Convergence in the Mediterranean fl oras in
central Chile and California: insights from comparative biogeography. In: Arroyo MTK, Zedler
PH, Fox MD (eds) Ecology and biogeography of Mediterranean ecosystems in Chile,
California, and Australia. Springer, New York, pp 43–88
Artigas J (1994) Entomología Económica: Insectos de interés agrícola, forestal, médico y veteri-
nario (nativos, introducidos y susceptibles de ser introducidos). Editorial Universidad de
Concepción, Concepción
Askevold IS (1991) On some poorly known, misidentifi ed and mislabelled Chilean Chrysomelidae
Coleoptera. Rev Chilena Entomol 19:11–15
Barriga TJE, Curkovic T, Fichet T et al (1993) Nuevos antecedentes de coleópteros xilófagos y
plantas hospederas en Chile, con una recopilación de citas previas. Rev Chilena Entomol
20:65–91
Beéche M, Sandoval A, Rothmann S et al (1999) Detección y control del gorgojo del eucalipto en
Chile, Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae). Servicio Agrícola y
Ganadero, Santiago
Beéche M, Goycoolea C, Rothman S et al (2003) Detección y control biológico de los taladradores
del eucalipto en Chile, Phoracantha semipunctata Fabricius y Phoracantha recurva Newman
(Coleoptera: Cerambycidae). Servicio Agrícola y Ganadero, Chile
S.A. Estay
393
Beèche M, Lanfranco D, Zapata M et al (2012) Surveillance and control of the Sirex woodwasp:
the Chilean experience. In: Slippers B, de Groot P, Wingfi eld MJ (eds) The Sirex woodwasp
and its fungal symbiont. Springer, Dordrecht, pp 229–245
Beggs J (2001) The ecological consequences of social wasps ( Vespula spp .) invading an ecosystem
that has an abundant carbohydrate resource. Biol Conserv 99:17–28
Burckhard D (1994) Generic key to the Chilean jumping plant-lice (Hemiptera: Psylloidea) with
inclusion of potential exotic pests. Rev Chilena Entomol 21:57–67
Burckhardt D, Elgueta M (2000) Blastopsylla occidentalis Taylor (Hemiptera: Psyllidae), a new
introduced eucalypt pest in Chile. Rev Chilena Entomol 26:57–61
Carrillo R, Cerda L (1987) Zoofi tófagos de Nothofagus Chilenos. Bosque 8:99–103
Carnegie AJ, Matsuki M, Haugen DA et al (2006) Predicting the potential distribution of Sirex
noctilio (Hymenoptera: Siricidae), a signifi cant exotic pest of Pinus plantations. Ann For Sci
63:119–128
Carnieletto C (2006) Ophelimus eucalypti Gahan (1922), (Hymenoptera: Eulophidae), la avispa
formadora de agallas. Chile For 327:47–48
Caron V, Ede F, Sunnucks P et al (2014) Distribution and rapid range expansion of the introduced
willow sawfl y Nematus oligospilus Förster (Hymenoptera: Tenthredinidae) in Australasia.
Aust Entomol 53:175–182
Cerda LA, Beéche MA (1989) Análisis general de las plagas insectiles asociadas a pino insigne y
eucalipto en Chile. Proceedings of the national symposium on forestry plant protection prob-
lems in Chile. IICA, Santiago
Chapman JD (1994) Notes on Mulanje cedar–Malawi’s national tree. Commonwealth For Rev
73:235–242, 272–273
Ciesla WM (1988) Pine bark beetles: a new pest management challenge for Chilean foresters. J For
86:27–31
Ciesla WM (1991) Cypress aphid, Cinara cupressi , a new pest of conifers in eastern and southern
Africa. FAO Plant Protect B 39:82–93
Ciesla WM (2003) European woodwasp: a potential threat to North America’s conifer forests.
J For 101:18–23
Curkovic T, Araya J, Guerrero M (2004) Avances en el manejo de la avispa chaqueta amarilla en
Chile. Aconex 84:19–24
Donoso C (1993) Bosques templados de Chile y Argentina. Variación, estructura y dinámica.
Editorial Universitaria, Santiago
Downer JA, Svihra P, Molinar RH et al (1988) New psyllid pest of California USA pepper tree.
Calif Agric 42:30–32
Eastop VF, Heie OE, Fuentes–Contreras E et al (1997) Notes on two new aphid species (Hemiptera:
Aphididae) detected in Chile. Rev Chilena Entomol 24:81–84
Edwards R (1976) The world distribution pattern of the German wasp, Paravespula germanica
(Hymenoptera: Vespidae). Entomol Ger 3:69–271
Elgueta M (2010) Gonipterus scutellatus el “gorgojo del eucalipto” (Coleoptera: Curculionidae)
en Chile. In: Lanfranco D, Ruiz C (eds) Entomología Forestal en Chile. Ediciones Universidad
Austral de Chile, Valdivia, pp 425–442
Elgueta M, Marvaldi AE (2006) Lista sistemática de las especies de Curculionoidea (Insecta:
Coleoptera) presentes en Chile, con su sinonímia. Boletín del Museo Nacional de Historia
Natural 55:113–153
Estay SA (2004) Insectos del arbolado urbano. Servicio Agrícola y Ganadero, Santiago
Estay SA, Lima M (2010) Combined effect of ENSO and SAM on the population dynamics of the
invasive yellow jacket wasp in central Chile. Popul Ecol 52:289–294
Estay SA, Araya JE, Guerrero MA (2002) Biología de Gonipterus scutellatus Gyllenhal
(Coleoptera: Curculionidae) en San Felipe, Chile. Boletín de Sanidad Vegetal Plagas
28:391–397
Estay SA, Navarrete SA, Rothmann SR (2012) Effects of human mediated disturbances on exotic
forest insect diversity in a Chilean Mediterranean ecosystem. Biodivers Conserv
21:3699–3710
13 Invasive Insects in the Mediterranean Forests of Chile
394
FAO (2008) Overview of forest pests, Chile. Forest Health & Biosecurity working papers
(FBS/12E). Forest Resources & Development Service, Rome
FAO (2009) Global review of forest pests and diseases. FAO forestry paper 156, Rome
Ferrada R, Ide S, Valenzuela J et al (2007) Intercepciones de insectos vivos realizadas en emba-
lajes de madera de internación en el período 1995–2005. Servicio Agrícola y Ganadero,
Santiago
Fuentes ER, Avilés R, Segura A (1990) The natural vegetation of a heavily man–transformed land-
scape: the savanna of central Chile. Interciencia 15:293–295
Fuentes ER, Miethke S, Avilés R (1993) Contemporary patterns of landscape change in areas with
a Mediterranean–type climate in central Chile. In: Turner BL, Gómez A (eds) Consequences of
the Columbian encounter. CSIC, España, pp 90–99
Fuentes–Contreras E, Munoz R, Niemeyer H (1997) Diversidad de áfi dos (Hemiptera: Aphidoidea)
en Chile. Rev Chil Hist Nat 70:531–542
González RH, Lamborot L (1989) El género Saissetia Deplanche en Chile:(Homoptera: Coccidae).
Acta Entomol Chil 15:237–241
González RH (1989) Insectos y ácaros de importancia agrícola y cuarentenaria en Chile. Edit.
Ograma, Santiago
González RH, Barría G, Guerrero (1986) Nematus desantisi Smith, a new species of forest impor-
tance in Chile (Hymenoptera: Tenthredinidae). Rev Chil Entomol 14:13–15
González P, Ide S, Jaques L (2010) Cinara cupressi Buckton (Hem: Aphididae), el pulgón del
ciprés. In: Lanfranco D, Ruiz C (eds) Entomología Forestal en Chile. Ediciones Universidad
Austral de Chile, Valdivia, pp 473–486
Goycoolea C, Beéche M, González P et al (2002) Detección y control del psílido de los eucaliptos
Ctenarytaina eucalypti (Hemiptera: Psyllidae). Servicio Agrícola y Ganadero, Santiago
Grez A, Zaviezo T, González G, Rothmann S (2010) Harmonia axyridis in Chile: a new threat.
Ciencia e Investigación Agraria 37:145–149
Grooves RH, Di Castri F (1991) Biogeography of Mediterranean invasions. Cambridge University
Press, Cambridge
Gutiérrez R (2011) Valor del arbolado urbano: la experiencia de la Municipalidad de Santiago.
Mundo For 21:4–6
Harris JWE, Wood RO (1967) The European pine shoot moth, Rhyacionia buoliana (Lepidoptera:
Olethreutidae), another introduced forest pest. J Entomol Soc BC 64:14–16
Hechenleitner V, Gardner MF, Thomas PI et al (2005) Plantas amenazadas del centro–sur de Chile.
Distribución, conservación y propagación. Universidad Austral de Chile y Royal Botanical
Garden of Edinburgh, Valdivia
Huerta A, Gómez IC, Puga K et al (2011) Life cycle of Xanthogaleruca luteola (Coleoptera:
Chrysomelidae) in Santiago, Chile, and sex fenotypo differentiation of adults. Boletín de sani-
dad vegetal Plagas 37:57–64
Ide S, Muñoz C, Beéche M et al (2006) Detección y control biológico de Glycaspis brimblecombei
Moore (Hemiptera: Psyllidae). Servicio Agrícola y Ganadero, Santiago
Ide S, Ruiz C, Sandoval A et al (2011) Detección de Thaumastocoris peregrinus (Hemiptera:
Thaumastocoridae) asociado a Eucalyptus spp . en Chile. Bosque 32:309–313
Ide S, Valenzuela J, Estay SA et al (2014) Presión de ingreso de insectos forestales exóticos a Chile
desde 1996. In: Jaksic F, Castro S (eds) Invasiones biológicas en Chile: causas globales e
impactos locales. Ediciones UC, Santiago, pp 437–457
INE (2014) Estadísticas demográfi cas y vitales.
http://www.ine.cl/ . Accessed 23 Jan 2015
INFOR (2008) Manejo integrado: técnica para la recuperación del crecimiento de Austrocedrus
chilensis . CORFO, Santiago
INFOR (2014) Estadísticas forestales chilenas 2014.
http://www.infor.cl . Accessed 23 Jan 2015
Kasper M (2004) The population ecology of an invasive social insect, Vespula Germanica
(Hymenoptera: Vespidae) in South Australia. Thesis, University of Adelaide, School of Earth
and Environmental Sciences, Discipline of Environmental Biology, Adelaide
S.A. Estay
395
Klein–Koch C, Waterhouse DF (2000) The distribution and importance of arthropods associated
with agriculture and forestry in Chile. Australian Centre for International Agricultural Research,
Canberra
Koch F, Smith DR (2000) Nematus oligospilus Förster (Hymenoptera: Tenthredinidae), an intro-
duced willow sawfl y in the Southern Hemisphere. Proc Entomol Soc Wash 102:292–300
Lanfranco D (2010a) Insectos defoliadores y sus efectos sobre sobre los sistemas forestales pre-
sentes en Chile. In: Lanfranco D, Ruiz C (eds) Entomología Forestal en Chile. Ediciones
Universidad Austral de Chile, Valdivia, pp 181–202
Lanfranco D (2010b) La polilla del brote del pino: como se enfrentó la primera y más importante
plaga forestal en Chile y sus implicancias actuales. In: Lanfranco D, Ruiz C (eds) Entomología
Forestal en Chile. Ediciones Universidad Austral de Chile, Valdivia, pp 409–424
Lanfranco D, Dungey HS (2001) Insect damage in Eucalyptus: a review of plantations in Chile.
Austral Ecol 26:477–481
Lanfranco D, Ide S, Ruiz C et al (2002) Agentes Entomopatógenos asociados a Productos
Forestales Primarios de Exportación. Editorial El Kultrún, Valdivia
Luebert F, Pliscoff P (2006) Synopsis of bioclimate and vegetation of Chile. Editorial Universitaria,
Santiago
Madden JL (1988) Sirex in Australasia. In: Berryman AA (ed) Dynamics of forest insect popula-
tions: patterns, causes, implications. Plenum Press, New York, pp 407–429
Mapondera TS, Burgess T, Matsuki M et al (2012) Identifi cation and molecular phylogenetics of
the cryptic species of the Gonipterus scutellatus complex (Coleoptera: Curculionidae:
Gonipterini). Aust J Entomol 5:175–188
Miller WE, Hastings AR, Wootten JF (1961) European pine shoot moth. Forest insect and disease
leafl et 59. USDA Forest Service, Ohio
Montalva C, Rojas E, Ruiz C et al (2010) El pulgón del ciprés en Chile: una revisión de la situación
actual y antecedentes del control biológico. Bosque 31:81–88
Muñoz R, Beéche M (1995) Antecedentes sobre dos especies de reciente identifi cación para Chile
(Homoptera: Aleyrodidae, Aphididae). Rev Chilena Entomol 22:89–91
Murphy ST, Nair KSS, Sharma JK (1996) Status and impact of invasive conifer aphid pests in
Africa. In: Varma RV (ed) Impact of diseases and insect pests in tropical forests. Proceedings
of the IUFRO Symposium, Peechi, pp 289–297
Olalquiaga FG (1952) El pulgón de los pinos en Chile. Rev For 16:91–92
Olivares TS (2000) Ctenarytaina eucalypti (MASKELL 1890): el psílido del eucalipto en Chile
(Hemiptera: Sternorryncha: Psylloidea: Spondyliaspininae). Gayana 64:239–241
Orondo SB, Day RK (1994) Cypress aphid ( Cinara cupressi ) damage to a cypress ( Cupressus
lusitanica ) stand in Kenya. Int J Pest Manag 40:141–144
Parra P (2010) La avispa taladradora de la madera de latifoliadas Tremex fuscicornis Fabricius
(Hymenoptera: Siricidae) en Chile. In: Lanfranco D, Ruiz C (eds) Entomología Forestal en
Chile. Ediciones Universidad Austral de Chile, Valdivia, pp 323–336
Peña MA, Altmann SH (2009) Use of satellite–derived hyperspectral indices to identify stress
symptoms in an Austrocedrus chilensis forest infested by the aphid Cinara cupressi . Int J Pest
Manag 55:197–206
Rodríguez F, Sáiz F (2014) Parasitoidism of Psyllaephagus pilosus Noyes (Hym.: Encyrtidae) on
the blue gum psyllid, Ctenarytaina eucalypti (Maskell) (Hem.: Psyllidae) in V Region euca-
lypts plantations. Agric Técn 66(4):342–351
Rojas E, Beèche M (2010) Detección y control de Sirex noctilio F. en Chile (Hymenoptera:
Siricidae). In: Lanfranco D, Ruiz C (eds) Entomología Forestal en Chile. Ediciones Universidad
Austral de Chile, Valdivia, pp 303–322
Rojas E, Gallardo R (2004) Manual de insectos asociados a maderas en la zona sur de Chile.
Servicio Agrícola y Ganadero, Santiago
Rundel PW (1981) The matorral zone of central Chile. In: di Castri F, Goodall DW, Specht RL
(eds) Mediterranean–type Shrublands. Elsevier, Amsterdam, pp 175–201
13 Invasive Insects in the Mediterranean Forests of Chile
396
Rundel PW (1998) Landscape disturbance in Mediterranean–type ecosystems: an overview. In:
Rundel PW, Montenegro G, Jaksic F (eds) Landscape disturbance and biodiversity in
Mediterranean–type ecosystems. Springer, Berlin, pp 3–22
Rundel PW, Montenegro G, Jaksic F (1998) Landscape disturbance and biodiversity in
Mediterranean–type ecosystems. Springer, Berlin
Ruz L (2002) Bee pollinators introduced to Chile: a review. In: Kevan P, Imperatriz Fonseca VL
(eds) Pollinating bees- the conservation link between agriculture and nature. Ministry if
Environment, Brazil, pp 155–167
SAG (2004) Informe anual 2004 subdepartamento de vigilancia y control de plagas forestales y
exóticas invasoras. Servicio Agrícola y Ganadero, Santiago
SAG (2005) Informativo fi tosanitario forestal N°2. Servicio Agrícola y Ganadero, Santiago
SAG (2006) Informativo fi tosanitario forestal N°3. Servicio Agrícola y Ganadero, Santiago
SAG (2010) Informativo fi tosanitario forestal N°4. Servicio Agrícola y Ganadero, Santiago
SAG (2014) Leptocybe invasa Fischer & Lasalle (Hymenoptera: Eulophidae) Microavispa forma-
dora de agallas en Eucalipto. Plagas y enfermedades.
http://www.sag.cl/ambitos–de–accion/
leptocybe–invasa–fischer–lasalle–hymenoptera–eulophidae–microavispa–formadora–de .
Accessed 23 Jan 2015
SAG (2015) Sirex noctilio o avispa de la madera del pino. Plagas y enfermedades.
http://www.sag.
cl/ambitos–de–accion/sirex-noctilio-o-avispa-de-la-madera-del-pino . Accessed 23 Jan 2015
San Martín J, Donoso C (1996) Estructura fl orística e impacto antrópico en el bosque Maulino de
Chile. In: Armesto JJ, Villagrán C, Arroyo MTK (eds) Ecología de los Bosques Nativos de
Chile. Editorial Universitaria, Santiago, pp 153–168
Schulz JJ, Cayuela L, Echeverria C et al (2010) Monitoring land cover change of the dryland forest
landscape of Central Chile (1975–2008). Appl Geogr 30:436–447
Silva SI, Bozinovic F, Jaksic FM (2005) Frugivory and seed dispersal by foxes in relation to mam-
malian prey abundance in a semiarid thornscrub. Austral Ecol 30:739–746
Stanković SS, Petrović A, Ilić Milośević M et al (2015) Morphological and molecular character-
ization of common European species Adialytus (Hymenoptera: Braconidae: Aphidiinae) based
on the mtCOI barcoding gene and geometric morphometrics of the forewings. Eur J Entomol
112:165–174
Valente C, Hodkinson I (2009) First record of the Red Gum Lerp Psyllid, Glycaspis brimblecombei
Moore (Hem.: Psyllidae), in Europe. J Appl Entomol 133:315–317
Villacide JM, Corley JC (2012) Ecology of the woodwasp Sirex noctilio : tackling the challenge of
successful pest management. Int J Pest Manag 58:249–256
Watson GW, Voegtlin DJ, Murphy ST et al (1999) Biogeography of the Cinara cupressi complex
(Hemiptera: Aphididae) on Cupressaceae, with description of a pest species introduced into
Africa. B Entomol Res 89:271–283
Zina V, Lima A, Caetano F et al (2012) First record of the pepper tree psyllid, Calophya schini
Tuthill (Hemiptera, Calophyidae), in the Palaearctic region. Phytoparasitica 40:127–130
S.A. Estay
