Trees, Forests and People 4 (2021) 100076

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Trees, Forests and People

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Flight activity and spread of Xylosandrus crassiusculus (Motschulsky)

(Coleoptera: Curculionidae) in Brazil
Luana de Souza Covre a, Adriano Arrué Melo b, Carlos Alberto Hector Flechtmann a,∗
a
Department of Plant Protection, FEIS/UNESP Ilha Solteira, São Paulo, Brazil
b
Department of Health Defense, Universidade Federal de Santa Maria, Rio Grande do Sul, Brazil

a r t i c l e i n f o a b s t r a c t

Keywords: The granulate ambrosia beetle (GAB) Xylosandrus crassiusculus is a polyphagous species with a worldwide distri-
Seasonality bution. The species is considered a serious pest in nursery, landscape and ornamental trees, and fruit orchards.
Voltinism The first specimens reported for Brazil were found along the coastal region in 2012. We report here the spread
Ethanol
inland of GAB based on ethanol-baited flight intercept trap surveys throughout the country. It is now present in
Ambrosia beetle
regions where orchards of several fruit tree species are planted, many of them known hosts of GAB. In this article,
Flight height
we also provide the first information on its flight activity in a subtropical climate in South America. In Barra do
Ribeiro, state of Rio Grande do Sul (southernmost state), GAB showed two flight peaks, both in the warmest
months of the year, while in Rio Branco, state of Acre (northern state), there were no clear peaks; these results
indicate that GAB is multivoltine in both localities. In vertical stratification studies in Tartarugalzinho, northern
state of Amapá, while GAB flew at up to 8 m high, flight was concentrated at ground level.

1. Introduction Premnobius cavipennis Eichhoff, 1878 and Scolytus rugulosus (Müller,

Man has been responsible for disseminating insect species into new
localities ever since crops and animals were introduced into new geo-
graphical areas (Morstatt 1941). However, the tremendous increase in
world trade in the last decades, coupled with difficulties in sampling
imported commodities, has led to a significant increase in the intro-
duction of exotic species (Campbell 2001; Work et al. 2005). Many
of these exotic species may threaten the native biodiversity by inter-
rupting ecological processes, while a number of them can become im-
portant pests (Vitousek et al. 1997; Mack et al. 2000; Roques 2010;
Liebhold et al. 2017).
Wooden articles pose a high risk for carrying invasive organisms,
mainly beetles (Campbell 2001; Haack 2006; Meurisse et al. 2019). Al-
though Brazil is among the largest world wood exporters (Bandara &
Vlosky 2012; ITTO 2019), it has not been spared from the introduction
of exotic beetle species. The main reason is that many of the imported
goods use boxes, pallets and other packaging material which is gener-
ally of low quality and that carries insects within it (Campbell 2001;
Haack 2006; Brockerhoff & Liebhold 2017).
Many non-indigenous ambrosia and bark beetles (Coleoptera: Cur-
culionidae: Scolytinae) have been introduced through commercial for-
eign trade in Brazil. The coffee berry borer, Hypothenemus hampei (Fer-
rari, 1867), the mango bark beetle, Cryphalus mangiferae Stebbing, 1914,
1818), for example, were “older introductions”- introduced in the first
half of the 20th century or even before (Wood 1982). However, a num-
ber of species were reported fairly recently, such as Amasa ‘truncata’
(Erichson, 1842), Cyrtogenius luteus (Blandford, 1894), and Xylosan-
drus crassiusculus (Motschulsky, 1866) (Flechtmann & Cognato 2011;
Flechtmann & Atkinson 2016; Flechtmann & Atkinson 2018).
The granulate ambrosia beetle (GAB), X. crassiusculus, was de-
scribed from Sri Lanka, southern Asia (Motschulsky 1866), and is
native to the Oriental region. It has spread worldwide, to North
(Anderson 1974; Douglas et al. 2013), Central (Kirkendall & Øde-
gaard 2007; Flechtmann & Atkinson 2016; EPPO, 2020) and South
America (Flechtmann & Atkinson 2016; Landi et al. 2017; Córdoba
& Atkinson 2018; Kirkendall 2018), Europe (Pennacchio et al. 2003;
Nageleisen et al. 2015; Gallego et al. 2017; Kavčič 2018), Africa
(Schedl 1962; Browne 1963; Wood & Bright 1992; EPPO, 2020;
Nel et al. 2020) and Oceania (Brockerhoff et al. 2003; EPPO,
2020).
Xylosandrus crassiusculus is a polyphagous species; more than 200
species of trees and shrubs, in over 60 families, are known as hosts
(Schedl 1962; Wood & Bright 1992). This species is considered a serious
pest in nursery, landscape, ornamental trees, young forest plantations,
and fruit orchards (Browne 1963; Mizell et al. 1994; Fulcher et al. 2012;
Ranger et al. 2016; EPPO, 2020).

∗
Corresponding author.
E-mail addresses: luanasouza.co@gmail.com (L.d.S. Covre), adrianoarrue@hotmail.com (A.A. Melo), carlos.flechtmann@unesp.br (C.A.H. Flechtmann).

https://doi.org/10.1016/j.tfp.2021.100076
Received 7 January 2021; Received in revised form 23 February 2021; Accepted 24 February 2021
Available online 27 February 2021
2666-7193/© 2021 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/)
L.d.S. Covre, A.A. Melo and C.A.H. Flechtmann
We report here the spread of X. crassiusculus in Brazil after its original
detection in coastal areas of the country, and provide the first informa-
tion on its flight activity in a subtropical climate in South America.
2. Materials and methods
2.1. New Xylosandrus crassiusculus records in Brazil
For over two decades one of the junior authors (CAHF, alone or
with collaborators) has been collecting bark and ambrosia beetles (Cur-
culionidae, Scolytinae) throughout Brazil, with ethanol-baited traps, in
various vegetation types. This ongoing study is entitled “Nationwide
Survey of Brazilian Bark and Ambrosia beetles (Coleoptera, Curculion-
idae, Scolytinae and Platypodinae)” (NWS for short). General method-
ology information regarding trap type, lure, trapping frequency and re-
lated information employed at the NWS are as described below. The
vast majority of the new records reported here originated from the
NWS.
We superimposed these new localities with maps showing the coun-
try distribution of (1) avocado (Persea americana Mill.) orchards and (2)
orchards of the temperate crops Japanese persimmon (Diospyros kaki
L.f.) and various species of Rosaceae (apple (Malus domestica Borkh.),
nectarine (Prunus persica (L.) var. nucipersica (Suckow.)), peach (Prunus
persica (L.)), pear (Pyrus communis L.) and plum (Prunus domestica L.)).
We chose these orchards based on reports from the literature that GAB
attacks these fruit tree species (Fraser 2014; CAB International, 2020;
DEFRA 2020; EPPO, 2020), and hence might become a pest of them in
Brazil.
2.2. Xylosandrus crassiusculus flight and seasonality
Study sites were a stand of Eucalyptus saligna Smith planted in May
2013 (30°22′56.46"S 51°8′6.15"W) and a semideciduous seasonal forest
fragment (30°25′45.37"S 51°8′28.84"W), both as part of the Horto Flore-
stal Barba Negra, owned by CMPC Celulose Riograndense, and located
in Barra do Ribeiro, state of Rio Grande do Sul, Brazil.
As established in the NWS protocol, in the center of each site we
deployed five flight intercept traps (modified from Berti Filho & Flecht-
mann 1986), placed at 1.5 m above ground and 25 m apart in a single
transect. Ethanol concentration was 96%, and its release rate was 870
mg/d at 20°C. The trapping frequency was weekly from 2 July 2018 un-
til 24 June 2019, equalling 52 trapping weeks. Voucher specimens are
deposited in the Museu de Entomologia da FEIS/UNESP (MEFEIS), Ilha
Solteira, state of São Paulo, Brazil.
Daily air temperature (maximum, minimum and mean), mean air
relative humidity and rainfall were obtained from the Meteorological
Station "Viveiro" (30°20′34.18"S 51°14′49.42"W), owned by CMPC, and
distant less than 14 km from the sampling sites.
In most regions of Brazil, the seasons can be classified into two peri-
ods, a “hot season”, with higher temperatures, normally associated with
more rainfall, and a “cold season”, with colder temperatures and usually
less precipitation (Flechtmann et al. 1995). Here we divided the collect-
ing period into a “hot season”, comprehending the period between 31
Aug 2018 until 18 Mar 2019, and the remaining period into a “cold sea-
son”. Meteorological values and GAB catches were compared between
“seasons” using generalized linear models (Proc GLM) and treatment
means were separated by the Tukey test ( SAS Institute, 1990).
Additional collecting sites from which data were here used and dis-
cussed were Aporé (state of Goiás), Garça (state of São Paulo), Paragomi-
nas, Santa Isabel do Pará (state of Pará), Pinheiro Machado (state of Rio
Grande do Sul), Rio Branco (state of Acre), Tartarugalzinho (state of
Amapá) and Venda Nova do Imigrante (state of Espírito Santo). Collect-
ing periods analized for each locality were Sep2018-Dec2019, Aug2017-
Aug2018, May2018, Aug2017-May2018, Oct2019-Aug2020, Apr2014-
Oct2018 and Oct2016-Nov2017, respectively.
2
Trees, Forests and People 4 (2021) 100076
3. Results
3.1. New records
ACRE: Rio Branco, Parque Zoobotânico da Universidade Federal do
Acre, bamboo-suppressed open ombrophilous forest (open terra firme
forest), 9°56′57.93"S 67°52′12.29"W, ethanol-baited FIT, 12-V-2020, 19-
VI-2020, 3-VII-2020, 5-VIII-2020, 14-III-2020, F.W.S. Silva (MEFEIS,
20); same except open ombrophilous forest (open terra firme forest)
with palms, 9°57′00.30′’S 67°52′45.14′’W, ethanol-baited FIT, 17-IV-
2020,30-VII-2020 (MEFEIS, 4).
AMAPÁ: Tartarugalzinho, Retiro Paraíba, terra firme om-
brophilous forest fragment, 1°07′30.5′’N 51°18′32.5′’W, 2-m high
ethanol-baited FIT, 12-XI-2016, 1-IV-2017, 4-VIII-2018, W.R. Silva
(MEFEIS, 3); same except 6-m high ethanol-baited FIT, 25-VII-
2015 (MEFEIS, 1); same except 8-m high, 24-XII-2016, 7-I-2017
(MEFEIS, 2).
ESPÍRITO SANTO: Venda Nova do Imigrante, Fazenda Alto Viçosa,
Pinus elliottii stand planted Sep 2000, 20°24′59.9"S 41°07′07.4"W,
ethanol-baited FIT, 30-XI-2016, 7-XII-2016, 15-II-2017, 21-IV-2017, 26-
IV-2017, 01-VI-2017, 7-VI-2017, 17-XI-2017, D.S. Martins (MEFEIS, 8).
GOIÁS: Aporé, Fazenda Barra, stand of a hybrid of Euca-
lyptus grandis x E. urophylla planted Feb 2011, 19°4′51.50"S
51°44′17.90"W, ethanol-baited FIT, 24-IX-2018, A.C. Menis
(MEFEIS, 3).
PARÁ: Paragominas, Mineração Paragominas - Norsk Hydro, post-
mining reforested area, 3°13′36.94"S 47°43′14.93"W, yellow tray, 8-V-
2018, 3-VI-2018, F.R.A. Almeida (MEFEIS, 2); Santa Isabel do Pará, So-
coco Agroindústrias da Amazônia, Cocos nucifera var. anão-verde stand
planted 2012, 1°13′14.62"S 48° 5′38.78"W, pitall trap with ethanol, IV-
2018, L.L.B. Carvalho (MEFEIS, 3); same except riparian forest fragment,
1°13′44.80"S 48°3′6.66"W, IV-2018 (MEFEIS, 1).
RIO GRANDE DO SUL: Barra do Ribeiro, CMPC Celulose Ri-
ograndense, Eucalyptus saligna stand planted May 2013, 30°22′56.46"S
51°8′6.15"W, ethanol-baited FIT, 17-VIII-2018, 21-VIII-2018, 24-IX-
2018, 1-X-2018, 12-XI-2018, 3-XII-2018, 2-I-2019, 4-II-2019, 15-
IV-2019 A.A. Melo (MEFEIS, 13); Pinheiro Machado, Fazenda
Guarda Velha, Olea europaea stand planted Oct2010, 31°29′58.70"S
53°30′31.50"W, ethanol-baited FIT, 14-II-2019, T. Scheunemann
(MEFEIS, 1).
SÃO PAULO: Garça, Fazenda Enseada, ex trunk of diseased/dying
Khaya senegalensis planted Mar 2014, 22°15′38.58"S 49°40′41.40"W, 07-
VI-2018, L.A. Benso (MEFEIS, 1).
Except for the record for São Paulo, all others were obtained through
the NWS study.
3.2. Flight and seasonal variation
Values for maximum (F1,49 = 19.74, P < 0.0001), minimum
(F1,49 = 18.37, P < 0.0001) and mean (F1,49 = 21.75, P < 0.0001) air
temperatures, plus air relative humidity (F1,49 = 23.42, P < 0.0001),
were significantly higher during the “hot season” when compared to
the “cold season”, while for rainfall there were no statistically dif-
ferences between “seasons” (F1,49 = 0.06, P = 0.8049). These results
legitimate the division of the trapping period into “hot” and “cold”
seasons.
GAB was trapped significantly more often in Barra do Ribeiro during
the “hot season” than during the “cold season” (F1,259 = 20.90, P <
0.0001). The first peak of GAB coincided with rising air temperatures
in mid-September. This small peak was followed by a much higher peak
that matched a period after rainfall receded, and lasted for five weeks,
from early November until the beginning of December. Outside these
two peak periods, trapping numbers were equal to or smaller than two
individuals per week, and no beetles were collected in the winter months
of May through July (Fig. 1).
L.d.S. Covre, A.A. Melo and C.A.H. Flechtmann
Fig. 1. Weekly values of mean temperature (maximum, average, minimum), relative humidity and rainfall, and sum of Xylosandrus crassiusculus catches in ethanol-
baited flight intercept traps in an Eucalyptus saligna stand. Barra do Ribeiro, Rio Grande do Sul state - Brazil, 2 July 2018 until 24 June 2019.
4. Discussion
The first records of GAB in Brazil were limited to the coastal area
of the country, near ports of entry (Flechtmann & Atkinson 2016). The
wide span among these reported localities suggest there might have been
multiple introductions (Fig. 2).
It is possible that GAB was introduced in Pará through Belém,
which has a large port, and from there spread ca. 40 km to Santa Is-
abel to Pará, and further south to Paragominas, over 200 km from the
coast.
It is clear that GAB spread quickly inland. In as little as five years, it
spread ca. 400 km into Espírito Santo, and distances exceeding 800 km
into Goiás, if Rio de Janeiro and São Paulo localities, respectively, are
used as initial reference points.
It is very difficult to ascertain the precise date a given species is in-
troduced into a specific region (Haack & Rabaglia 2013). However, for
Amapá we are certain that GAB reached Tartarugalzinho only in 2015,
3
Trees, Forests and People 4 (2021) 100076
when we collected the first specimen, because we were trapping inten-
sively in this locality with the same trap and lure since April 2014. Be-
cause this region is characterized as sparsely inhabited and dominated
by the Amazon rainforest, both its natural and human-aided spread
would be expected to be slow. Hence, it is possible that GAB was es-
tablished in Macapá, only 120 km from Tartarugalzinho, at an earlier
date than 2014, when it was first reported there (Flechtmann & Atkin-
son 2016).
In Barra do Ribeiro, it seems that GAB is well established - 177 spec-
imens were trapped in a stand of Eucalyptus saligna. It might well be
that this beetle spread from Rivera in Uruguay, where it was reported
in 2013 (Landi et al. 2017), to Rio Grande do Sul - Rivera is distant ca.
400 km from Barra do Ribeiro and only 180 km from Pinheiro Machado.
We are not sure how X. crassiusculus spread to Acre. This locality
is within and surrounded by the Amazon forest, and possibly it was
introduced from neighboring Peru, even though the species was not yet
reported in that country (Smith et al. 2017).
L.d.S. Covre, A.A. Melo and C.A.H. Flechtmann
Even though GAB is able to colonize natural forests (Maetô
et al. 1999; Kirkendall & Ødegaard 2007; Ulyshen & Sheehan 2019;
Sanguansub et al. 2020), as shown here in Acre and Amapá (Fig. 2), this
was not proved to be the case in Aporé and in Barra do Ribeiro. While
in those localities GAB was trapped in eucalypt stands, no specimens
were trapped in the riparian forest and cerrado (savannah-like vegeta-
tion) fragments in Aporé (C.A.H. Flechtmann et al., unpublished data),
and in the nearby semideciduous seasonal forest fragment in Barra do
Ribeiro. There is no good explanation for this fact, especially consid-
ering that GAB is a highly polyphagous species (Schedl 1962; Wood &
Bright 1992; EPPO, 2020).
GAB was already reported attacking several Eucalyptus species
(International 2020; DEFRA 2020; EPPO, 2020), which might explain
trappings of this species in eucalypt stands in Aporé and Barra do
Ribeiro. Even though it would be possible that GAB was developing in
4
Trees, Forests and People 4 (2021) 100076
Fig. 2. Occurrence of Xylosandrus crassiusculus
and distribution of area planted with avocado
orchards (Persea americana Mill.) in Brazil.
shrubby species in the eucalypt understory instead, if this were the case,
then it should have been trapped in the native surrounding vegetation
in both of these stands.
We report for the first time GAB attacking the trunk of a dis-
eased/dying Khaya senegalensis (Desr.) tree. The species has already
been reported attacking several Meliaceae species, including Khaya
grandifoliola C. DC. in Nigeria (Roberts 1969) and Khaya ivorensis A.
Chev. in Ghana (Browne 1963).
It is worth noting that GAB already occurs in both the N-S latitu-
dinal and E-W longitudinal frontiers of Brazil. This is a clear indica-
tion that this ambrosia beetle has faced no climatic local constraints,
so it is expected to spread through the vast majority of the coun-
try. Considering the speed of the spread, based on data from their
initial reports (Flechtmann & Atkinson 2016), we believe that within
20 to 30 years GAB should be distributed throughout the country.
L.d.S. Covre, A.A. Melo and C.A.H. Flechtmann
Fig. 3. Occurrence of Xylosandrus crassiusculus and distribution of area planted with the temperate crops Japanese persimmon (Diospyros kaki L.f.) plus Rosaceae
orchards in Brazil.
Perhaps only the semi-arid region of northeastern Brazil, also known
as caatinga, and characterized by extremely dry conditions (Daly &
Mitchell 2000), could pose a challenge to colonization by this invading
species.
The currently known distribution of GAB matches some of the lo-
calities where avocado (Fig. 2) and persimmon + rosaceous fruit tree
orchards (Fig. 3) are planted in Brazil. Considering GAB will likely be
present in the majority of the country, a larger number of already (and
potentially new) host plants will be exposed to colonization by the bee-
tle, and it has the potential to become a pest in a few or even several
fruit orchard species and ornamental trees, as indicated in the literature
(see “Introduction”).
There are a few factors that could have influenced the efficacy of
detecting GAB in Brazil, including the attractant used and its release
rate. Ethanol was used as a lure in our experiments. From the liter-
ature, it is clear that GAB is attracted to this kairomone. There must
be an optimal – and yet unknown - release rate which is most at-
tractive to this species. However, it seems that ethanol is attractive
to GAB over a surprisingly wide range of release rates. In some ar-
ticles, the reported release rates were on the lower part of the spec-
trum (7 – 16 mg/d) (Werle et al. 2012; Burbano and Wright, 2012),
some others can be placed at intermediate levels (65 – 300 mg/d)
(Reding et al. 2013; Miller et al. 2018; Rabaglia et al. 2020), while the
majority were on the higher end of the range (500–1140 mg/d) (Oliver &
Mannion 2001; Miller & Rabaglia 2009; LaBonte 2010; Coyle et al. 2015;
Miller et al. 2020). We avoided here using the term ‘ultra high re-
lease rate’ due to the fact it is loosely used, covering a range from
300 mg/d to 1000 mg/d (Oliver & Mannion 2001; LaBonte 2010;
Rabaglia et al. 2020). Perhaps the authors were influenced by how the
company where they purchase the lures classify them. The release rate
in our experiments was at the higher end of the spectrum, which might
have helped in avoiding under-collecting GAB when present in small
numbers in a particular locality.
5
Trees, Forests and People 4 (2021) 100076
Another factor that might have influenced trapping would be the
height of the trap above ground. We set our traps at 1.5 m above ground
in our NWS sites; from experience, we know that in Brazil most Scolyti-
nae species fly somewhat close to the ground. From the literature, even
though GAB is known to be able to fly up to 20 m high (Maetô et al. 1999;
Miller et al. 2020), it is more frequently trapped close to the ground,
from 0.3 m to 0.8 m (Maetô et al. 1999; Miller et al. 2020; Ulyshen &
Sheehan 2019) or even up to 1.7 m (Reding et al. 2010). In Macapá, state
of Amapá, we set traps every two meters from ground level up to 18 m,
in a vertical stratification experiment (C.A.H Flechtmann et al., unpub-
lished data). GAB was trapped up to 8-m high traps (same maximum
height where we trapped GAB in Tartarugalzinho, Amapá). However,
58% of the specimens were trapped in ground level traps, and another
23% of them in 2-m high traps. These results match with the literature,
and it seems that our 1.5-m high traps in our NWS experiments were in
the predominant range of GAB flight height.
Our classification of the collecting period in Barra do Ribeiro into
two seasons matched long series of recorded temperature and rainfall
data found in the literature (Britto et al. 2008; Diniz et al. 2018). Con-
sidering that rainfall is fairly homogeneous throughout the year, it is
reasonable to assume that temperature was the main factor involved in
observed numbers of trapped GAB (Fig. 1). It seems that GAB behaves
in this locality the same way as in southern United States and in Italy,
where peaks in flight activity are temperature-influenced and are ob-
served in warmer months (Cote 2008; Labonte 2010; Reding et al. 2010;
Reding et al. 2013; Rassati et al. 2016; Williams & Ginzel 2019).
We had ethanol-baited traps out in the field also in Rio Branco, state
of Acre, northern Brazil, in the Amazon forest domain. In this local-
ity average monthly, average temperatures never drop below 23°C and
it rains throughout the whole year (Duarte 2006). We trapped GAB
specimens from March through August 2020, but no clear peak was
observed (C.A.H Flechtmann et al., unpublished data). This might in-
dicate that GAB faces no temperature and rainfall constraints in this
L.d.S. Covre, A.A. Melo and C.A.H. Flechtmann
region, and hence could likely reproduce continuously in northern
Brazil.
Our trapping results suggest that GAB is multivoltine, in both the
southernmost state of Rio Grande do Sul (with milder temperatures)
and especially in the northern state of Acre (with warmer tempera-
tures). These results match reports from the literature, that show that
this species is multivoltine in Asia (Wu et al. 1978) and in various
United States regions (Oliver & Mannion 2001, Reding et al. 2010;
Werle et al. 2012; Williams & Ginzel 2019).
Based on our findings, we were able to verify that GAB moved deeply
inland from their original coastal region localities, and it is now present
in all regions of Brazil. GAB has already reached several fruit orchard
regions, where it might inflict economic damage, among other possible
host plants of economic or ornamental value. In its continuing spread,
GAB is expected to establish itself throughout the country, due to minor
or no climatic restrictions at all. The damage potential of GAB will likely
be enhanced in Brazil due to the fact it can generate several generations
per year. While it is able to fly up to 8 m high, most flights concentrate
below 1 m height. Hence, traps should be placed within this height in-
terval in monitoring programs, where ethanol should be used as a lure.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
CAHF would like to thank the NWS partners Wilson R. Silva
(Amapá), Farley W.S. Silva (Acre), David S. Martins (Espírito Santo),
Aline C. Menis (Goiás), Lana L.B. Carvalho and Roberta M. Valente
(Pará), Tiago Scheunemann and Dori E. Nava (Rio Grande do Sul) and
Lucas A. Benso (São Paulo) for assistance with the trappings, André R.
Farias (EMBRAPA Territorial) for putting together and providing the
shapefiles, to Silvia Y. Tanabe and Nathalia M.L. Silva for assistance
in compiling GAB data and to Roger A. Beaver for invaluable help in
reviewing this manuscript.
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