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M.Orlova-Bienkowskaja, M.Volkovitsh. 2017. ... A case study of the native range of the EAB...

М.Я. Орлова-Беньковская

M.J.Orlova-Bienkowskaja, M.G.Volkovitsh. 2017.
Are native ranges of the most destructive invasive pests well known? A case study of the native range of the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae).
[М.Я. Орлова-Беньковская, М.Г. Волкович. Хорошо ли мы знаем ареалы опасных инвазивных вредителей? Исследование естественного ареала ясеневой изумрудной узкотелой златки (ЯИУЗ) Agrilus planipennis (Coleoptera: Buprestidae).]
Biological Invasions. 123.
DOI: 10.1007/s10530-017-1626-7.
Published on-line 15.11.2017.
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Are native ranges of the most destructive invasive pests well
known? A case study of the native range of the emerald ash
borer, Agrilus planipennis (Coleoptera: Buprestidae)

Marina J. Orlova-Bienkowskaja, Mark G. Volkovitsh

Received: 25 June 2017 / Accepted: 13 November 2017

Springer International Publishing AG, part of Springer Nature, 2017

Knowledge of the native range of invasive
pests is vital for understanding their biology, for
ecological niche modeling to infer potential invasive
distribution, and for searching of natural enemies.
Standard descriptions of pest ranges frequently pass
from one publication to another without verification.
Our goal is to test the reliability of distributional
information exemplified by the native range of one of
the most destructive and most studied invasive forest
insect pests of Asian origin: the emerald ash borer
(EAB), Agrilus planipennis Fairmaire. Since the first
detections of this notorious insect pest in North
America in 2002 and European Russia in 2003, it
has killed hundreds of millions of ash trees. Based on
the examination of museum specimens and literature
sources we compiled the most comprehensive data-
base of records (108 localities) and the most detailed
map of the native range of EAB in East Asia to date.
There are documented records for 87 mainland
localities of EAB in the Russian Far East (Primorskiy,
Khabarovskiy Kray), China (Heilongjiang, Jilin,
Liaoning, Beijing, Hebei, Tianjin, Shandong and
Xinjiang), and South Korea, and 21 localities in Japan.
Records from Nei Mongol, Sichuan, Mongolia, and
Taiwan are ambiguous since no documented records
are available. The example of EAB shows that
standard descriptions of pest ranges could include
false or ambiguous data. Compilation of the database
of documented localities is the only way to obtain
reliable inform ation on the range.

Keywords: EAB, Pest, Ash, Fraxinus, East Asia, Native range

Knowledge of the native ranges of invasive pests is of
great importance. First, distributional information is
crucial for understanding the various aspects of the
biology and ecology of the pest. Second, natural
enemies of the pest occur throughout its native range
and these may be used for biological cont rol. Third,
the precise information about the native range, and
especially the database of document ed localities, form
the basis for ecological models and forecasts of the
potential invasive distribution of the pest (Sobek-
Electronic supplementary material The online version of
this article ( con-
tains supplementary material, which is available to authorized

M. J. Orlova-Bienkowskaja (&)
A.N. Severtsov Institute of Ecology and Evolution,
Russian Academy of Sciences, 33 Leninskiy Prospect,
Moscow, Russia 119071
M. G. Volkovitsh
Zoological Institute, Russian Academy of Sciences,
Universitetskaya nab. 1, St., Petersburg, Russia 199034

   Publications on certain invasive
species usually include a ''standard'' description of its
distribution in the terms of administrative units: 
countries and provinces in which the species is
recorded. These range descriptions frequently pass
from one publication to another without critical
verificat ion. Our goal is to test the reliability of
distributional information based on a re-examination
of the native range of one of the most destructive and
the most studied forest pest s in the world: Agrilus
planipennis Fairmaire, 1888.
The emerald ash borer (EAB), Agrilus planipennis
native to East Asia is an extremely aggressive invasive
insect pest of ash (Fraxinus spp.) (Jendek and
Grebennikov 2011; Cha morro et al. 2015). It was first
detected in North America in 2002 (Haack et al. 2002)
and in European Russia in 2003 (Izhevskii 2007;
Volkovitsh 2007) and since then it has killed hundreds
of millions of ash trees in both regions (Haack et al.
2015). The economic and environmental impact of the
invasion is tremendous (Herms and McCullough
2014). The standard descriptio n of the native range
is as follows: China: Beijing, Hebei, Heilongjiang,
Jilin, Liaoning, Nei Mongol, Shandong, Sichuan,
Tianjin, Xinjiang; Japan : Hokkaido, Honshu, Kyushu,
Shikoku; North Korea; South Kor ea; Taiwan; Mon-
golia; Russia: Far East (Primorskyi Kray) (Hou Tao-
qian 1986; Jendek and Grebennikov 2011; Chamorro
et al. 2015; EPPO 2017). Samples of distributio n
maps, based on administrative units, can be found in
Chamorro et al. (2015) and Haack et al. (2015).
In the invaded North American range, more than
5000 occurrence records are known (Sobek-Swant
et al. 2012), and range expansion is tracked by the
website Emerald Ash Borer Info (2017). The EAB
invaded range in European Russia is also rather well
known with more than 70 occurrence records docu-
mented (Straw et al. 2013; Orlova-Bienkowskaja
2013, 2014). However, only limited information is
available on the occurrence of EAB in its native range
because it is considered a rar e species in natural forests
(Alexeev 1979; Yurchenko et al. 2007). Distributional
information is scattered in many publications, some of
them are in Russian, Chinese and Japanese or
published in rare, early sources not available on the
internet. Distribution maps of EAB withi n its native
range are presented in several papers, but all of them
are incomplete since they are based on a relatively
small number of precise localities. Liu et al. (2003)
and Wei et al. (2004) compiled distribution maps of
EAB in China based on 11 and 14 documented
localities, respectively. Keever et al. (2013) mapped 8
localities in China and South Korea. Bray et al. (2011)
mapped 18 localities in China, South Korea and Japan.
Sobek-Swant et al. (2012) adopted localities from
Bray et al. (2011), together with some additional
information, and compiled a map based on 36
localities in mentioned countries and the Russian Far
East. It is the most comprehensive map of EAB
occurrence to date.
Very early information on occurrence of EAB in
some regions is not reliable because many adminis-
trative borders have change d since time of publication.
The problem is complicated by a number of unsolved
taxonomic questions and the fact that in the southern
parts of the range , A. planipennis could be confused
with closely related species, particularly with A.
tomentipennis Jendek and Chamorro recorded from
Laos and Taiwan (Jendek and Chamorro 2012;
Chamorro et al. 2015). To verify the EAB distribution
in East Asia, we collected all the information available
about EAB occurrence records (totaling, 108 points)
and re-examined its native range, taking into account
these unresolved issues.
Agrilus planipen nis was described from China
(Fairmaire 1888). The following names are now
regarded as synonyms of A. planipennis: Agrilus
marcopoli Obenberger, 1930, which type locality is
unclear, Agrilus feretrius Obenberger, 1936 described
from Taiwan and Agrilus marcopoli ulmi Kurosawa,
1956 described from Japan (Chamorro et al. 2015).
When analyzing the native range of EAB, it is
important to consider the host plants and biotopes
(native vs. anthropogenic ) in which the p est is
recorded in specific locations. Under natural condi-
tions, EAB larval feeding is reported mainly on native
Fraxinus mandshurica Rupr.; F. chinensis ssp. rhyn-
chophylla (Hance) A. E. Murray (including F. japon-
ica Blume ex K. Koch as its synonym); and
occasionally on F. chinensis chinensis Roxb.
Throughout its invasive range, as well as in anthro-
pogenic landscapes within its native range, it is
recorded on American ash species (F. americana L.;
F. nigra Marshall; F. pennsylvanica Marshall; F.
quadrangulata Mic hx.; F. velutina Torr.) and Euro-
pean ash (F. excelsior L. ) (Jendek and Grebennikov
2011; Jendek and Pola?kova 2014; Orlova-Bienkows-
kaja 2014; Chamorro et al. 2015). Within the invasive
range in the USA., larval feeding on another repre-
sentative of Oleaceae, white fringetree, Chionanthus
virginicus L. was recently reported (Cipollini 2015).
In Japan A. planipennis has been also reported from
Juglans mandshurica var. sie boldiana Maxim.; Pte-
rocarya rhoifolia Siebold and Zucc. (both Juglan-
daceae); and Ulmus davidiana var. japonica (Rehder)
Nakai (Ulmaceae) (Jendek and Grebennikov 2011;
Jendek and Pola?kova 2014), however there is no
evidence whether EAB was reared from or only
collected on these plants. The only verified larval hosts
of EAB are Fraxinus species (Zhao et al. 2005; Jendek
and Pola?kova 2014) and Chionanthus virginicus
(Cipollini 2015), both from the family Oleaceae.
Native hosts and biotopes are most valuable to outline
the native range of EAB, while the regions where the
pest occur only on introduced hosts in anthropogenic
landscapes, could represent a secondary range.

We compiled a data base comprising 87 localities of
Agrilus planipennis in mainland East Asia and 21
localities in Japan , totaling 108 localities (see elec-
tronic supplementary material) and made a detailed
map of its native range using the software DIVA-GIS
7.5 (Fig. 1). The sources of information were mainly
the specimens deposited in the collections of the
Zoological Institute of Russian Academy of Sciences
(ZIN; Russia, St. Petersburg), National Museum
(NMPC, Prague, Czech Republic) (type specimens
of A. marcopoli Obenberger, 1930 and A feretrius
Obenberger, 1936); M.Yu. Kalashian (MKCY; Arme-
nia, Yerevan) and S.N. Ivanov (SICV; Russia, Vladi-
vostok), as well as the exact localities from 31
publications (see Table 1). Some localities are still
known only from old labels, though man y geograph-
ical names, administrative divisions, and borders have
partly changed since the date of specim en collection.
Therefore, a significant part of the work was the
interpretation of old geographical labels.
To obtain additional information on the EAB
localities in the Russian Far Ea st, we requested the
curators of the largest Russian Coleoptera collections
to check for the presence of specimens of EAB in their
corresponding collections. There were no specimens
in the collections of Zoological Museum of Moscow
State University (ZMUM; A.A. Gusakov, personal
communication), Moscow State Pedagogical Univer-
sity (MSPU; K.V. Makarov, personal communica-
tion), and Siberian Zoological Museum (Coleoptera
collection of Siberian Zoological Muse um 2017).
Only 8 specimens of EAB collected by G.I. Yurch-
enko on Fraxinus pennsylvanica in Khabarovsk in
2008 were found in the collection of the Federal
Scientific Centre of Biodiversity of Terrestrial Biota of
East Asia, Far East Department of Russian Academy
of Sciences (FSCB; Russia, Vladivostok) (S.A. Sha-
balin, personal communication).

Documented records
Records based on exact localities of A. planipennis are
listed in the Table 1; occurrence points are docu-
mented in the database (see electronic supplementary
material) and mapped in Figs. 1, 2, 3.

Russian Far East
Prior to the beginning of this century, EAB was
exceptionally rare in the forests of the Russian Far
East, as evidenced by its rarity in most Coleoptera
collections (see ''Methods'' section). Under natural
conditions it feeds on native ash species Fraxinus
mandshurica and F. chinensis rhynchophylla, but it
has never been considered a pest, since it affects only
weakened trees (Yurchenko et al. 2007). No outbreaks
in the natural ash stands have been reported (Yurch-
enko 2016). The outbreaks have been recorded in
urban plantations on Fraxinus pennsylvanica only
recently. Cultivation of F. pennsylvanica in the Far
East started in the early 1950s. At first, plantations of
this ash species existed in the arboretum of Gorno-
taezhnaya station of Academy of Sciences of the
USSR (Primorskiy Kray) (Flora USSR 1952) and in
the Far East Forestry Research Institute arboretum
(Khabarovskiy Kray) (Yurchenko 2009). Then, in the
1950s-1980s, F. pennsylvanica was widely cultivated
in urban plantations in Far East cities. Currently, the
plantations of F. pennsylvanica in Vladivostok,
Khabarovsk and othe r cities of Primorskiy and
Khabarovskiy Krays are severely damaged by EAB
(Yurchenko 2016). All these points of occurrence are
shown in Fig. 2 (green and red circles in Primorskiy
and Khabarovski y Krays).
Agrilus planipennis was first collected in the Far
East in 1935 (Yurchenko et al. 2007) but first recorded
for the Russian fauna in 1979 (as A. marcopoli
Obenberger) (Alexeev 1979). Only a few specimens
were collected in the Far East before 2004 in six
districts of Primorskiy Kray: Hasan, Spassk, Shko-
tovo, Lazo, Ternei and Chuguevka Districts (Alexeev
1979; Jendek 1994; Yurchenko et al. 2007; Jendek and
Grebennikov 2011 ; examined specimens from ZIN,
MKCY, SICV). Then, in 2004-2012, special surveys
revealed EAB damaging introduced F. pennsylvanica
all over Primorskiy Kray and also in the south of
Khabarovskiy Kray (Yurchenko 2009 ; Belokobylskij
et al. 2012; Duan et al. 2012; Yurchenko et al.
2007, 2010, 2013a, b).
In 2010 and 2011 the survey of plantations of ash
trees in Sakhalin Island did not reveal the presence of
EAB (Yurchenko 2016). It also has not been found in
Kuril Islands (M.Yu. Proschalykin, personal

In China, as in the Russian Far East, EAB does not
pose a serious threat to natural ash stands or planta-
tions attacking mainly ornamental trees in urban
landscapes. Documented records are known from
Heilongjiang, Jilin, Liaoning, Beijing, Hebei, Tianjin,
Shandong and Xinjiang (see Table 1, Fig. 1 and
electronic supplementary material). Localities of sev-
ere outbreaks are indicated in the map in Fig. 2).
In Heilongjiang A. planipennis was a very rare
species until the 1960s. The first outbreak was
recorded in Harbin in 1964 on Fraxinus americana
introduced from America (Wei et al. 2004). The
damage to plantati on of Northeast Forestry University
Experimental Forest was so severe that the entire
plantation was removed. In the cours e of surveys in
2003-2008, EAB was recorded in four prefectures of
Heilongjiang on its native host plants Fraxinus
mandshuric a and F. chinensis rhynchophylla (Liu
et al. 2003; Wei et al. 2004; Bray et al. 2011; Keever
et al. 2013). Jendek and Grebennikov (2011) indicated
that a specimen with the label ''Primorsk 13.8.1935
Fig. 1 Range of EAB in East Asia. 1, 2, 3: native range, type
locality and other documented localities of A. planipennis in
mainland Asia; 4: ambiguous type locality of A. marcopoli;
5: region in Xinjiang, where A. planipennis was discovered in
2016; 6, 7: ambiguous records of A. planipennis; 8 : the range
of A. planipennis in Japan; 9: type locality of A. marcopoli ulmi
(synonym of A. planipennis); 10 : other documented localities
of A. planipennis in Japan; Kh: Khabarovskiy Kray, P: 
Primorskiy Kray, Hei: Heilongjiang, J: Jilin, L: Liaoning,
B: Beijing, Heb: Hebei, T: Tianjin, Sh: Shandong, Xi: 
Xinjiang, Si: Sichuan, NM: Nei Mongol. Sources of infor-
mation are listed in Table 1

Table 1 Documented records of Agrilus planipennis in East Asia
Region Number of mapped localities
Years of collection
Host plants Sources of information
Russia: Primorskiy Kray
31 1935-2012 F. mandshurica, F. chinensis
rhynchophylla, F. pennsylvanica*
Specimens in ZIN, MKCY, SICV,
Alexeev (1979), Jendek (1994),
Yurchenko et al. (2013a, b, 2007),
Volkovitsh (2009), Jendek and
Grebennikov (2011), Belokobylskij
et al. (2012), Duan et al. (2012)
Russia: Khabarovskiy Kray
6 2004-2011 F. mandshurica, F. pennsylvanica* 
Specimens in FSCB, Yurchenko et al.
(2007), Yurchenko (2009), Duan et al.
(2012), Sobek-Swant et al. (2012)
China: Heilongjang
6 1964-2008 F. mandshurica, F. chinensis
rhynchophylla, F. americana*
Yu (1992), Liu et al. (2003), Wei et al.
(2004), Bray et al. (2011), Keever et al.
(2013), Wang et al. (2016 )
China: Jilin 10 1986-2008 F. mandshurica, F. chinensis
rhynchophylla, F. pennsylvanica*
Hou Tao-qian (1986), Yu (1992), Liu
et al. (2003, 2007), Wei et al.
(2004, 2007), Bray et al. (2007, 2011),
Wang et al. (2016)
China: Liaoning 10 1986-2013 F. mandshurica, F. chinensis
rhynchophylla, F. chinensis chinensis, F.
Hou Tao-qian (1986), Yu (1992), Wei
et al. (2004), Liu et al. (2003, 2007),
Bray et al. (2011), Wang et al. (2016)
China: Beijing 3 1888-2013 F. velutina* 
Fairmaire (1888), Jendek (1994), Zhang
et al. (2005), Bray et al. (2011), Wang
et al. (2016)
China: Hebei 5 1937-2008 F. chinensis rhynchophylla, F. chinensis
chinensis, F. velutina*
Jendek (1994), Liu et al. (2003), Wei
et al. (2004), Bray et al. (2011)
China: Tianjin 4 1982-2008 F. chinensis rhynchophylla, F. chinensis
chinensis, F. velutina*, F. pennsylvanica*
Liu et al. (2003), Wei et al. (2004, 2007),
Bray et al. (2007, 2011)
China: Shandong
2 Pre 2003 F. velutina* Yu (1992), Wei et al. (2004)
China: Xinjiang 0 Before 2003 and 2016
Wei et al. ( 2004), Zhaozhi Lu (personal
South Korea 10 1943-2008 F. chinensis rhynchophylla Kurosawa (1956), Bray (2007, 2011),
Belokobylskij et al. (2012), Keever
et al. (2013), GBIF (2017)
Japan: Hokkaido
4 1941-1974 Ulmus propinqua (= Ulmus divadiana var.
japonica), Pterocarya rhoifolia, Juglans
mandshurica var. sieboldiana, Fraxinus
mandshurica var. japonica
Kurosawa (1956), GBIF (2017),
Chamorro et al. (2015)
Japan: Honshu 14 1943-2003 Kurosawa (1956), Jendek (1994),
Akiyama and Akiyama (1996),
Schaefer (2004), GBIF (2017),
Chamorro et al. (2015)
Japan: Kyushu 2 1949-1979 Kurosawa (1956), Chamorro et al.
Japan: Shikoku 1 Pre 1956 Kurosawa (1956), Schaefer (2004)
Introduced ash species are marked with an asterisk
Mai He [river]'' was collected in Heilongjiang, but
actually Mai He river (currently, Artiomovka river) is
located in Primorskiy Kray, Russia. EAB is recorded
in at least five prefectures of Jilin province on F.
mandshurica, F. chinensis rhynchophylla, and F.
pennsylvanica, and at least in six prefectures of
Liaoning province on F. mandshurica, F. chinensis
rhynchophylla, and F. chinensis chin ensis (Hou Tao-
qian 1986; Wei et al. 2004; Liu et al. 2003, 2007 ; Bray
et al. 2011).
In Beijing Agrilus planipennis is known from the
late 19th century, since it is a type locality of the
species (Fairmaire 1888). Now it is rather common in
this region and in the adjacent provinces of Hebei and
Tianjin on native Fraxinus chinensis rhynchophylla ,
F. chinensis chinensis, and the non-native (introduced
from North America) F. velutina, though before the
introduction of the later it was rather rare there (Li
Zhong 2002; Liu et al. 2003; Bray et al. 2011; Jendek
and Grebennikov 2011; Chamorro et al. 2015 ).
Fraxinus velutina was imported to Tianjin in 1952
and to Beijing in 1956 (Wei et al. 2004). In Tianjin city
EAB was first recorded in 1982 on a plantation of F.
velutina (Wei et al. 2004). The efforts to control the
outbreak failed and all infested trees were cut down in
1991. In 1993 F. velutina was planted in Guangang
District of Tianjin and severe damage by EAB was
observed in 1998-2002. During surveys in
2003-2008, A. planipennis was found in at least six
localities in Tianjin on F. velutina and F. pennsylvan-
ica (Liu et al. 2003; Wei et al. 2004; Bray et al.
2007, 2011). The outbreaks were also recorded in
Hebei (Wei et al. 2004).
Fig. 2 Outbreaks of Agrilus planipennis within its native range.
1: outbreak points (1964: outbreak on Fraxinus americana in
Harbin; 1982 and 1998: outbreaks on F. velutina in Tian-
jin; * 2000: outbreaks on F. pennsylvanica in Vladivostok
and Khabarovsk); 2, 3: range and points of occurrence of A.
planipennis in mainland Asia; 4, 5: range and points of
occurrence of A. planipennis (= A. marcopoli ulmi) in Japan.
Sources of information are listed in Table 1
In the province of Shandong, the southernmost
documented EAB localities in mainland China, it
occurs on F. velutina in urban plantations (Wei et al.
2004). Based on the introduced host, it may be
suggested that localities in Shandong belong to the
introduced range after EAB's primary range expan-
sion followed the outbreaks in Tianjin in 1982 and
1998 (Fig. 2) (Fuester and Schaefer 2006).
EAB is also recorded in the north-west part of
China in Xinjiang Autonomous region (Wei et al.
2004). According to Zhaozhi Lu (personal communi-
cation), in 2016, EAB was found in Yili valley. The
distribution map of F. mandshurica (Fig. 3) shows
that this ash species can reach the westernmost limits
of Xinjiang. Additionally, another ash species, F.
angustifolia syriaca (Boiss) Yalt. (= F. sogdiana
Bunge), occurs in western Xinjiang and adja cent
regions of Kazakhstan (Flora of China 2017). The
recent detection of emerald ash borer in Xinjiang
should be treated as a new invasion by the pest, rather
an expansion of the pest's native range.

Korean peninsula
EAB is known from Korea at least since 1943
(Kurosawa 1956). Special surveys made in
2003-2008 have shown that EAB is widespread there
but occurs at low densities and feeds mainly on
Fraxinus chinensis rhynchophylla and F. mandshurica
(Ko Je Ho 1969; Williams et al. 2006, 2010; Bray et al.
2007, 2011; Keever et al. 2013). All documented
localities are from South Korea, but there is no doubt
that the species also occurs in North Korea because
this region is situated in the center of the known range
of the species (EPPO 2017).

Records of EAB from Japan are based on the
assumption that Agrilus marcopoli ulmi is a synonym
of A. planipennis (Table 1, Figs. 1, 2). In 1956 A.
marcopoli ulmi, with the host plant Ulmus propinqua
(= davidiana var. japonica according to Jendek and
Fig. 3 Ranges of the native host plants of Agrilus planipennis
in East Asia. 1: Fraxinus mandshurica, 2: F. mand-
shurica & F. chinensis (F. chinensis chinensis and F. chinensis
rhynchophylla), 3: F. chinensis, 4: points of occurrence of A.
planipennis in mainland Asia, 5: points of occurrence of A.
planipennis in Japan. Sources of information on ranges of
Fraxinus species: Cleary et al. (2016), GBIF (2017). Sources of
information for EAB points of occurrence are listed in Table 1
Pola?kova 2014), was described from Sapporo, Japan
based of specimens collected in 1930 (Kurosawa
1956). Jendek (1994) synonymized A. marcopoli ulmi
with A. planipennis, but Akiyama and Ohmomo
(1997) treated it as a distinct subspecies and indicated
that its host plants are Ulmus divadiana var. japonica;
Pterocarya rhoifolia; Juglans mandshurica var. sie-
boldiana (= ailanthifolia according to Jendek and
Pola?kova 2014); Fraxinus mandshurica var. japonica.
However, it is unclear if the specimens examined by
these authors were reared from the men tioned plants or
just collected on them. Bray et al. (2011) established
that the mtDNA haplotype from a sing le adult spec-
imen collected in Japan strongly differed from all the
A. planipennis collected in Asia and North America.
Thus, the presence of the emerald ash borer in Japan
has been doubted by some authors (Herms and
McCullough 2014). However, the identity of A.
planipennis and A. marcopoli ulmi was currently
confirmed by E. Jendek who claims that there are no
reliable morphological features allowing the separa-
tion of mainland and Japanese specimens even at a
subspecific level (E. Jendek, personal communication
of June 2017).

Ambiguous records

The occurrence of EAB in Mongolia is cited in
hundreds of publications (e.g. Ko Je Ho 1969; Hou
Tao-qian 1986; Jendek 1994; Li Zhong 2002; Jendek
2006; Jendek and Grebennikov 2011; Dua n et al.
2012; Chamorro et al. 2015). But all these records
actually refer to a single locality, namely the type
locality of Agrilus marcopoli Obenberger, 1930,
which is the junior synonym of A. planipennis (Jendek
1994). The lectotype of A. marcopoli bears the label
''Mongol. or.: Chan-heou'' (lectotype: NMPC). It is
difficult to determine where this geogr aphical point is
situated. According to the catalogue of Carabus
collecting localities in China, ''Chan-heou'' is situated
in Hebei (China) (40.38 N, 117.41 E) (Schufitze, Kle-
infeld 2007). According to E. Jendek (personal
communication), the name ''Chan-heou'' refers to
the settlement Cha-hen-hua (currently Ujimqin) on
Shiliin (Xilin) Gol river (46.63 N, 119.52 E), which is
situated in Nei Mongol (China) near the Mongolian
border (Googl e maps 2017). Therefore, the exact
position of the type locality of A. marcopoli is not
clear, but some authors believe that it is situated in
China rather than in Mongoli a (e.g. Schaefer 2004). 
In 2003, Schaefer traveled to Mongolia looking for A.
planipennis but did not find it. Mongolian entomolo-
gists with whom he has consulted, said that EAB does
not occur in this country (Schaefer 2004). Addition-
ally, neither the genus Fraxin us, nor the family
Oleaceae are reported from Mongolia (Grubov
1955, 1982; Urgamal 2014). Therefore, according to
the information available to date, there is no reason to
believe that EAB occurs in Mongolia.

Nei Mongol (Inner Mongolia) (China)
Nei Mongol is regarded to be a part of the native range
of EAB in many sources (Hou Tao-qian 1986; Yu
1992; Jendek 1994; Nonnaisab et al. 1999; Li Zhong
2002; Wei et al. 2004; Jendek and Grebennikov 2011;
Chamorro et al. 2015), but no documented localities
are available. Jendek and Grebennikov (2011) indi-
cated that two specimens labelled ''Chahar Yangki-
aping 8-10.VII.1937'' were collected in Nei Mongol.
But monastery Yangkiaping is situated in the histor-
ical region Chahar, currently in Hebei (Yan et al.
2013). Additionally, it is not excluded that the type
locality of Agrilus marcopoli, ''Chan-heou'', is really
situated in Nei Mongol (see above). However, the
special survey in Nei Mongol in 2006 did not reveal
this species in the region (Wang et al. 2010 ). The host
plant Fraxinus mandshurica has very restricted dis-
tribution in Nei Mongol (Cleary et al. 2016), but
occurs just in the region, which is supposed to be the
type locality of A. marcopoli (Fig. 3).

Sichuan (China)
The record of EAB from Sichuan is based on a single
old specimen from the National Museum in Prague
(Czech Republic) labelled ''Szechuan'' without any
additional information (Jendek 1994). Th e origin of
this specimen is unknown, and it cannot be excluded
that this geogr aphic label is incorrect. According to
Wei et al. (2004), the host plant Fraxinus chinensis,
occurs in this provi nce, but EAB was not found nether
in Sichuan nor in adjacent provinces of China. This
region is situated more than 1000 km south-west from
the nearest documented locality of EAB.

There are only two old records of EAB from Taiwan.
Firstly, Agrilus feretrius Obenber ger, 1936 (lectotype:
NMPC), which is a junior synonym of A. planipennis
(Jendek 1994 ), was described from ''Formosa'' (old
name of Taiwan). Secondly, Schaefer (2004) exam-
ined specimen(s) of A. feretrius (as A. teretrius) from
Taoyuan Province of Taiwan in the collection of
Kurosaw a (National Museum, Tokyo) and concluded
that the later treated A. feretrius as a subspecies of A.
marcopoli (= A. planipennis) thoug h he has never
validated this status. Herms and McCullough (2014)
doubted the presence of EAB in Taiwan, since the
records could refer to another species and we agree
with this point of view. First, Taiwan is situated far to
the south of all othe r known loca lities of EAB. Second,
there are at least two closely related species from the
same species group reported from Taiwan: A. tomen-
tipennis Jendek and Chamorro 2012 and A. pseudol-
ubopetri Jendek and Chamorro 2012 (Chamorro et al.
2015). Additionally, though two species of Fraxinus
are widely distri buted in Taiwan (Flora of Taiwan
1998), these are not indicated as host-plants for EAB.

Agrilus planipennis was reported from Laos (Jendek
and Grebennikov 2011). However, later it was estab-
lished that this record referred to the closely related A.
tomentipennis from the same species-group (Jendek
and Chamorro 2012; Chamorro et al. 2015).

Careful re-examination of the native range of A.
planipennia h as shown, that the reliable range is much
more restricted than it is usually described in lite ra-
ture. There is no doubt that A. planipennis occurs in the
Far East of Russia (Khabarovskiy Kray, Primorskiy
Kray); China (Heilongjiang, Jilin, Liaoning, Beijing,
Hebei, Tianjin, Shandong, Xinjiang), and the Korean
peninsula. Each of these records is confirmed by
several independent sources of information, occur-
rence records are well documented and the identifica-
tion of the species is relia ble. From a biogeographical
viewpoint, the bulk of the native range of EAB is
within the Stenopean (Ma nchurian-Northern-
Chinese-Northern-Japanese) nemoral region and
areas transitional to adjacent regions of the Palaearctic
part of East Asia (Emeljanov 1974; Konstantinov et al.
2009). Over a great extent, it also overlaps with the
eastern part of the range of the manchurian ash
(Fraxinus mandshurica Rupr.) (Fig. 3), which is one
of the principal EAB host plants. Thus, EAB can be
characterized as a mostly nemoral species. This is also
confirmed by the location and extent of invasive
ranges of EAB in North America and European Russia
(see Chamorro et al. 2015). At the same time, taking
into account the anthropogenic factor (the presence of
urban plantations in populated areas) and the possi-
bility of spreading ash trees along the river valleys
(which can explain some remote isolated locations of
the pest), the EAB range may extend far beyond the
nemoral zone (e.g., location in Xinjiang) forming
satellite colonies at the periphery of the main range as
it is observed, for example, in its invasive area in
European Russia (Orlova-Bienkowskaja 2013 , 2014)
and North America (Siegert et al. 2014).
In this regard, it is unknown if the entire modern
range of A. planipennis in East Asia is native (primary
range) or it has partly expanded as a result of transition
of the pest to the cultivated American ashes (secondary
range). Because humans plant trees wherever they
choose, pests often take advantage of new host sources,
ranging far outside their ancestral home to invade the
new plantings (Fuester and Schaefer 2006). Such cases
are rather common among coleopteran species. For
example, Agrilus convexicollis expanded its range into
European Russia because of the mass cultivation of
alien Fraxinus pennsylvanica and because of drastic
weakening of its plantations as a result of the invasion
of EAB (Orlova-Bienkowskaja and Volkovitsh 2014).
It can be assumed that the transition from resistant
native to non-resistant introduced American ash
species (most commonly F. pennsylvanica) triggered
the pest outbreaks (Fig. 2) and led to a significant
expansion of the primary range (Khabarovsk in Russia,
Shandong and, possibly, Xinjiang in China). Moreover,
this transition followed by outbreaks can be regarded as
an initial phase of subsequent invasion of the EAB
beyond East Asia.
It seems that just the outbreaks of EAB in Tianjin
and Hebei were the initial point of further invasion of
the pest to North America and European Russia. First,
EAB has been a rare species in native forests, so the
probability of invasion of the pest from regions other
than regions of outbreaks is vanishingly small, since
high propagule pressure is necessary to overwhelm the
ecological resistance of the community (Holle and
Simberloff 2005). Second, the time of the outbreaks in
China coincides with the estimated time of invasion: it
is assumed that EAB was unintentionally introduced
to North America and European Russia in the early
1990s (Siegert et al. 2014; Straw et al. 2013). Third,
genetic researc h by Bray et al. (2011) has revealed that
North American populations are most closely related
to populations from the Chines e province of Hebei and
Tianjin City. Moreover, this region is an important
center of Chinese export, where goods from this region
are exported all over the world. Unintentional intro-
duction with exported goods could be the vector of
EAB invasion.

EAB is one of the most popular research subjects in
the fields of forest entomology, invasion biology, plant
protection, urban ecology, etc. over the past 15 years.
According to Google Scholar, about 100 scientific
publications dealing with this notorious pest are
published each year. But, despite the great interest to
EAB after its invasion of North America and European
Russia, its native range was poorly known. The majority
of the publications that mention EAB distribution
include unverified information from earlier sources.
False or ambiguous records of species distribution
sometimes occur because of misidentifications, taxo-
nomic problems and unclear labels result in incorrect
interpretations of geographic data. The same concerns
arise with some other invasive organisms (Orlova-
Bienkowskaja et al.
2015). The example of EAB shows
that standard descriptions of native ranges of invasive
pests, repeated in hundreds of books and articles, could
be unreliable. Careful examination of range based on
collection of data on precise locations is the only way to
compile a reliable description. are still unresolved.
Acknowledgements The study was supported by Russian
Science Foundation, Project No 16-14-10031. We would like to
thank E. Jendek (Bratislava, Slovakia) for the valuable
information and comments on EAB distribution and providing
us with missing literature; we are also grateful to 
V. Kuba`n (S?lapanice near Brno, Czech Republic), G. I. Yurchenko (The
Far East Forest Research Institute, Forest Management Agency
of the Russian Federation, Khabarovsk, Russia), Zhaozhi Lu
(Xinjiang Institute of Ecology and Geography, Chinese
Academy of Sciences), and M. Yu. Proschalykin (FSCB) for
valuable information used in this paper; to M. Yu. Kalashian
(Institute of Zoology, Scientific Center of Zoology and
Hydroecology, National Academy of Sciences of Armenia,
Yerevan, Armenia), and S. N. Ivanov (private collector,
Vladivostok, Russia) for providing the exact locations for EAB
specimens in their collections for this study; to A. A. Gusakov
(ZMUM), K. V. Makarov (MSPU), and S. A. Shabalin (FSCB)
for checking the presence of EAB specimens in corresponding


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