Environmental Entomology, XX(XX), 2020, 1–9

doi: 10.1093/ee/nvaa147
Population Ecology Research

Long-Distance Migration of the Globe Skimmer Dragonfly

to Japan Revealed Using Stable Hydrogen (δ 2H) Isotopes
Keith A. Hobson,1,2,6 Hiroshi Jinguji,3 Yuta Ichikawa,4 Jackson W. Kusack,1 and

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R. Charles Anderson5

1
Department of Biology, University of Western Ontario, 1151 Richmond Street, London, ON N6A 5B7, Canada, 2Environment and
Climate Change Canada, Saskatoon, SK, Canada, 3Faculty of Food and Agricultural Sciences, Fukushima University, Fukushima,
Japan, 4Okazaki Senior High School, Aichi, Japan, 5Manta Marine, P.O. Box 2074, Malé, Republic of Maldives, and 6Corresponding
author, e-mail: khobson6@uwo.ca

Subject Editor: Rodrigo Mercader

Received 31 July 2020; Editorial decision 16 October 2020

Abstract
The globe skimmer dragonfly, Pantala flavescens Fabricius (Odonata: Libellulidae), is a long-distance migrant, well
adapted to exploiting ephemeral waterbodies. This species occurs in Japan every summer, but overwintering has
only been recorded on subtropical Ishigaki Island. It is not known from where the summer immigrants originate, nor
what proportion of the globe skimmers seen in Japan are of local origin. We analyzed stable hydrogen isotope (δ2H)
composition of wings of 189 P. flavescens captured at six sites in Japan from August to September in 2016 (n = 57)
and from April to November in 2017 (n = 132). We determined that the majority of individuals were immigrants.
Individuals of probable Japanese origin occurred only later in the year and were of lower mass on average than
immigrants. Immigrants potentially originated from a broad area as far west as northern India and the Tibetan
Plateau and, especially late in the season, as near as northcentral China and the Korean peninsula. However, for
April samples, the most parsimonious interpretation suggested southern origins, in northern Myanmar to southern
China, or possibly Borneo-Sulawesi. Our investigation underlines the power of combining stable isotope data
with other information such as wind speed and direction, arrival dates, and body mass to estimate origins and to
understand the life history of this and other insects.

Key words: Dragonfly migration, stable isotope, deuterium, migratory connectivity

One of the most spectacular examples of animal migration is the
intercontinental movement of the globe skimmer dragonfly Pantala
flavescens Fabricius. As its common name implies, this species is
broadly distributed across the globe (Fraser 1936, Russell et al. 1998,
Corbet 1999) and one of its most prominent hypothesized move-
ments involves the multigenerational circular trip from northern
India, potentially including the Himalayas and the Tibetan plateau,
through east Africa, on a journey exceeding 14,000 km (Fraser
1936, Anderson 2009, Hobson et al. 2012a). These movements are
facilitated in part by a short larval stage (~6 wk, compared with a
more typical 10 mo or more), which allows the species to make use
of ephemeral (often monsoonal) waterbodies, relatively free from
long-lived predators (Corbet 1999). Migratory flights also make use
of high-altitude winds (especially those associated with movements
of the Inter-Tropical Convergence Zone, ITCZ) and benefit from the
use of gliding combined with powered flight, as well as an ability to
feed on the wing while traveling (Anderson 2009, Li et al. 2014).
Despite widespread interest in the origins and movements
of P. flavescens, little is known about its migratory behavior and
current reports are largely anecdotal. This lack of understanding de-
rives from the fact that it is extremely difficult to track individual
small insects, especially at large spatial scales, since technological
advances in tracking vertebrates have typically not been refined for
insects (Chapman et al. 2015, but see Wikelski et al. 2006, Knight
et al. 2019). However, the advent of stable isotope-tracking methods,
which rely solely on endogenous signals present in animals, has revo-
lutionized the way in which origins of small insects can be tracked
(Hobson and Wassenaar 2019). The most prominent use of this tech-
nique has been directed at the migration of the eastern population
of the Monarch Butterfly, Danaus plexippus Linnaeus (Lepidoptera:
Nymphalidae), primarily using stable isotope measurements of deu-
terium (2H) in wing tissue (Wassenaar and Hobson 1998, Hobson
et al. 1999). This approach has also been used by other researchers
interested in Monarch patterns of spring recolonization in North
America (Miller et al. 2012, Flockhart et al. 2013), the effects of
natal origin on parasite loads (Altizer et al. 2015), the role of wing
coloration in flight distance (Hanley et al. 2013), and conservation
concerns related to individual origins (Flockhart et al. 2017). Stable

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2 Environmental Entomology, 2020, Vol. XX, No. XX

isotope tracking of other insect species such as dragonflies (Hobson
et al. 2012a) including P. flavescens (Hobson et al. 2012b, Cao et al.
2018) have since been conducted, and there is considerable promise
that this approach will help elucidate origins of insect migrants on
continental scales (reviewed by Stefanescu et al. 2016, Hobson et al.
2018, Hallworth et al. 2018). Such broad-scale analysis is required
since work evaluating the population genetic structure of P. fla-
vescens has revealed gene flow across large geographic areas, sug-
gesting near-panmixia (Troast et al. 2016, Alvial et al. 2019; but see
Pfeiler and Markow 2017).
In Japan, P. flavescens is a well-known summer migrant.
passed (Asahina and Turuoka 1967, 1968, 1969, 1970; Hashimoto
and Asahina 1969).
Our objective here was to delineate possible origins of P. fla-
vescens captured at various locations in Japan in 2016 and 2017
using stable hydrogen isotope analysis of wings (Hobson et al.
2012a, b). Captures took place during April–November, and we
were especially interested in separating migrants from locally bred
dragonflies. Prevailing winds during the summer are largely from
mainland Asia toward Japan, and we speculated that nonlocal P. fla-
vescens originated from East Asia, possibly from as far as northern
India and Mongolia.

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Throughout most of the country, it occurs in large numbers during
June–September, although some may be seen as early as April and as
late as November (e.g., Kuwada 1972, Sugiura et al. 1999, Tanaka
and Higashi 2003, Arai 2007, Iwata et al. 2009). Temperatures are
mostly too cold for it to survive over winter, with eggs failing to
hatch below about 15°C (Iwata et al. 2009, Ichikawa et al. 2016).
However, in the subtropical southern islands, temperatures are po-
tentially warm enough for P. flavescens to survive year-round and
successful overwintering of larvae has been recorded on Ishigaki
Island (Iwata et al. 2009, Watanabe 2019). Nevertheless, it is be-
lieved that most P. flavescens migrate into southern Japan every
spring from warmer countries to the southwest and that from there
the population rapidly increases, spreading northwards as summer
progresses. For example, Arai (2007) reported that many adult P. fla-
vescens were observed in the Ryukyu (Okinawa) Islands in spring
and then moved northward as far as Hokkaido. Pantala flavescens
is well known for its opportunistic reproductive strategies, with fe-
males laying eggs in small ponds and rice paddy fields, as well as
swimming pools and puddles (Watanabe 1999, Sato 2004, Arai
2012, Hirake 2012). Females can lay more than 1,000 eggs per day
if they encounter suitable oviposition sites (Ichikawa and Watanabe
2014). Eggs and larvae develop rapidly, completing development
within roughly 50 d in Japan (Iwata et al. 2009, Ichikawa et al.
2016) as they do elsewhere (Hawking and Ingram 1994, Suhling
et al. 2004). These biological characteristics result in rapid local
population increase, with large numbers from what are believed to
be locally bred subsequent generations being seen throughout Japan
during the summer and autumn (Wakana 1959, Kuwada 1972,
Arai 2007, Iwata et al. 2009). It is also known that typhoons in-
fluence the local distribution and abundance of P. flavescens, e.g.,
with dragonflies being observed from June to October on a weather
ship located off southern Japan in the Pacific Ocean (29°N, 135°E),
but with particularly large numbers being recorded when typhoons
Materials and Methods
Field Collections
Our field collections are summarized in Table 1. Pantala flavescens
specimens were individually stored at ambient condition for 24–48 h
in paper envelopes. Following excretion of feces, body mass was
measured using an electronic balance (to 0.0001 g). The sex and
stage of individuals were identified by wing condition and color ac-
cording to Ichikawa and Watanabe (2014). Specimens were then
frozen and stored dry in paper envelopes until shipment to the
Stable Isotope Hydrology and Ecology Laboratory of Environment
Canada. All P. flavescens were individually numbered and their sex,
maturity, and body weight were recorded (Fig. 1).
Stable Isotope Analyses
Following rinsing in 2:1 chloroform:methanol and overnight drying
in a fume hood, samples of 0.35 ± 0.02 mg of wing membrane were
weighed into pressed silver 3.5 × 5 mm capsules and analyzed using
a Eurovector Uniprep autosampler (Milan, Italy) carousel attached
to a Eurovector 3000 Elemental Analyzer, coupled with a Thermo
Delta V Plus isotope ratio mass spectrometer (Bremen, Germany) in
continuous flow mode with the helium carrier gas. After the samples
were loaded, the Uniprep autosampler (heated to 60°C) was vacuum
evacuated and subsequently flushed with dry helium twice to re-
move adsorbed atmospheric moisture from the crushed silver cap-
sules. The autosampler was then held under positive helium pressure
for the duration of the analytical run. Two USGS keratin standards,
EC-01 (formerly CBS: Caribou Hoof Standard) and EC-02 (KHS:
Kudu Horn Standard of Environment Canada), were included every
10 samples. An internal laboratory standard, powdered keratin (MP
Biomedicals Inc., Cat No. 90211, Lot No. 9966H) was included
to monitor instrument drift and provide a check on accuracy over

Table 1. Summary of stable hydrogen isotope values of wings (δ 2Hw) of individual Pantala flavescens analyzed in this study

Sample Date Latitude longitude δ 2Hp (‰) δ 2Hw (‰) n Statistics

2016
Miyagi 11 Aug. 38°13′ N 140°49′ E −57 −146.1 ± 10.8 20 a
Saitama 25 Aug. 35°47′ N 139°28′ E −56 −138.4 ± 8.0 11 ab
Shiga 7 Sept. 35°15′ N 136°13′ E −58 −131.6 ± 14.1 20 b
Hokkaido 15 Sept. 43°04′ N 141°20′ E −54 −145.7 ± 9.5 6 a
2017
Kagoshima 9–30 Apr. 31°21′ −31°44′ N 131°00′ −130°15′E −45 −110.5 ± 8.9 34 a
Kagoshima 30 May–9 Jun. 31°19′ N 130°54′E −45 −142.1 ± 9.1 6 b
Shizuoka 24 July−5 Aug. 34°44′ −35°01′ N 138°04′ −138°23′E −55 −130.9 ± 22.7 12 c
Kagoshima 5 Oct.−12 Nov. 31°16′ −31°23′ N 131°04′ −130°55′E −45 −129.3 ± 8.2 40 c
Shizuoka 24 Oct.–13 Nov. 34°39′ N 137°46′ −137°52′E −55 −123.9 ± 10.1 40 c

The δ 2Hp values correspond to predicted mean annual precipitation. δ 2H values were calculated for these locations using the waterisotopes.org calculator. The
letters in the statistics column indicate differences among groups as determined by Tukey’s post hoc (P < 0.05) tests.
Environmental Entomology, 2020, Vol. XX, No. XX
Fig. 1. Prefecture areas of collection of Pantala flavescens in Japan 2016–2017
(Global Administrative Areas 2018).
the course of each analytical session. Samples were combusted at
1,350°C using glassy carbon. Values of δ 2H of nonexchangeable
hydrogen were derived using the comparative equilibration ap-
proach of Wassenaar and Hobson (2003) and calibrated to VSMOW
using EC-01 (±1.9‰ 1 SD, n = 18, accepted δ 2H = −197.0‰) and
EC-02 (±1.6‰, n = 17, accepted δ 2H = −54.1‰). We used these
former calibration standard values because our original assignment
algorithms were based on these (Hobson et al. 1999). This approach
results in identical assignments if new calibration standards are used
with correspondingly modified assignment algorithms (see below;
Soto et al. 2017). In addition, because our original calibrations
linking precipitation δ 2H with dragonfly wing δ 2H were based on
combustion in a high-temperature glassy carbon reactor, we elected
to continue with this method versus combustion at lower temperat-
ures using a chromium-based reactor (Gehr et al. 2015, see also Soto
et al. 2017). Overall measurement error for EC-01, EC-02, and our
internal commercial powdered keratin δ 2H was ~2‰.
Assignment to Origin
We depicted the origins of wild dragonflies caught in Japan using
a likelihood-based assignment method (Hobson et al. 2009, Van
Wilgenburg et al. 2012, Wunder 2010), using the wing chitin δ 2H
isoscape (δ 2Hw) and an amount-weighted precipitation-to-wing cali-
bration algorithm derived by Hobson et al. (2012a). This was used
to convert amount-weighted mean annual precipitation δ 2H (δ 2Hp)
isoscapes (Bowen et al. 2005, Bowen 2019) into a δ 2Hw isoscape
(Fig. 2). We used the 14.7‰ residual SD error from this regression
in our assignments (Hobson et al. 2012b). We then created a spatial
layer representing the known geographic range for P. flavescens in
south and east Asia including India and used this as a mask (i.e., clip)
to limit our assignment to origin analyses.
We estimated the likelihood that a cell (pixel) within the δ 2Hw
isoscape represented a potential origin for a sample by using a
normal probability density function to estimate the likelihood func-
tion based on the observed δ 2Hw and thus depicted the likely origins
3
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Fig. 2. Expected wing δ 2H isoscape for Pantala flavescens in southeast Asia.
of each dragonfly by assigning individuals to the δ 2Hw isoscape one
at a time. We arbitrarily chose a 2:1 odds ratio to include only those
pixels (coded 1) with at least a 67% probability of origin versus all
others (coded 0). This resulted in a map of binary values for each
assigned individual representing presence (1) or absence (0) for each
cell in the isoscape. We then summed the results of individual assign-
ments by stacking the surfaces. We made geographic assignments to
origin using functions within the R statistical computing environ-
ment (R Development Core Team 2019) using scripts employing the
‘raster’ v. 3.0–12 (Hijmans 2020), ‘maptools’ v. 1.0-1 (Bivand and
Lewin-Koh 2020), and ‘rnaturalearth’ v 0.1.0 (South 2017) pack-
ages. Thus, the final assignment surface depicted the number of indi-
viduals coassigned at each pixel based on the odds criteria.
We derived an expected value for Japanese-origin P. flavescens by
averaging the mean annual (amount-weighted) precipitation δ 2Hp
for all of our Japanese sites (−57‰) based on the waterisotopes.org
calculator. That resulted in a predicted mean δ 2Hw value for P. fla-
vescens of −94.4‰ based on the algorithm derived by Hobson et al.
(2012b). We then allowed an SD of 14.7‰ to give a corresponding
range of −109.1 to −79.7‰ to be consistent with possible origins
from Japan.
Trajectory Analysis
We explored possible origins and immigration routes of our three
larger samples (n > 30) with the HYSPLIT (HYbrid Single Particle
Lagrangian Integrated Trajectory) model developed by the U.S.
National Oceanic and Atmospheric Administration (NOAA) Air
Resources Laboratory (Stein et al. 2015), using the GDSA1 dataset
to generate backwards trajectories from our sample locations (www.
ready.noaa.gov/hypub-bin/trajasrc.pl). We used arbitrary 5 d (120 h)
backward trajectories, with starting altitudes of 500, 1,000, and
15,000 m, and then in some cases investigated further with add-
itional starting altitudes at 250-m intervals. Arrival dates during the
first half of each collecting period in Japan were investigated. There
was no attempt to systematically generate large numbers of trajec-
tories in order, e.g., to estimate probabilities of origin from different
sectors. Rather, the aim was simply to demonstrate that at least some
trajectories linked potential areas of origin (as demonstrated by
our δ 2Hw analyses) with our sampling sites. Representative trajec-
tory plots are illustrated in Supp Material (online only; in two cases,
trimmed to 4 d [96 h] for clarity of presentation).
4
Fig. 3. Distribution through time of wing δ 2H values measured for Pantala
flavescens collected at four sites in Japan in 2016: Miyagi—11 August,
Saitama—25 August, Shiga—7 September, Hokkaido—15 September.
Also indicated is the mean and SD of the expected value for this species if
produced in Japan.
Fig. 4. Probable origins of Pantala flavescens collected in 2016 at (a) Miyagi prefecture—11 August, (b) Saitama—25 August, (c) Shiga—7 September, and (d)
Hokkaido—15 September. The legend indicates the number of individuals assigned to a given pixel consistent with the 2:1 odds criterion chosen (see Materials
and Methods).
Environmental Entomology, 2020, Vol. XX, No. XX
Results
For 2016 collections (11 August–15 September), there was a signifi-
cant difference in δ 2Hw among the four collection sites (Fig. 3; F3,
53
= 5.82, P = 0.02; Table 1). No clear changes in δ 2Hw were ob-
served across sites throughout the sampling period, although almost
all individuals were more depleted in 2H than expected for Japan.
We depicted origins of P. flavescens from each of these sites and col-
lection periods (Fig. 4) and found the highest probability of possible
origins in August and September were from outside Japan and cen-
tered in northern China or Mongolia and northern India. Few of our
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2016 samples appeared likely to have originated in Japan itself, with
just a small number of individuals collected in Shiga Prefecture on
7 September having δ 2Hw levels compatible with being locally bred
(Fig. 3).
For 2017 collections (9 April–13 November), there was a
significant difference in δ 2Hw among the five collection groups
(Fig. 5; F4, 127 = 20.52, P = 0.000) with three groups identified
(Kagoshima in April, Kagoshima in May vs all other samplings;
Table 1). The most enriched group was sampled at Kagoshima
Environmental Entomology, 2020, Vol. XX, No. XX
in April, but all samples except that collected in Kagoshima in
May–June had a proportion of individuals that could have poten-
tially been produced in Japan (Table 1; Fig. 5). We depicted ori-
gins of P. flavescens from each site and collection period in 2017
in Fig. 6. Our April sample from Kagoshima indicated possible
origins in Japan as well as sites as far as northern China and also
to the southwest in the region of northern Myanmar to southern
China, or Borneo-Sulawesi. Those more southwestern origins
were more plausible due to the dry season throughout much of
the far northwestern portion of the range under consideration and
our examination of wind trajectories in April (Supp Fig. S1 [on-
line only]). Later season captures in 2017 indicated possible ori-
gins throughout the range potentially including Japan.
Based on our derivation of expected values for Japanese-produced
individuals, it was possible to examine characteristics of individuals
potentially derived from Japan and those that were clearly immi-
grants. Potential Japanese-origin dragonflies were of lower mass
than immigrants (Table 2, Wilcoxon rank-sum test, P < 0.001), but
we found no difference in the proportion of males and females or
adults and immatures among clear immigrant and possible Japanese-
produced individuals (Fisher exact test, P > 0.05).
Discussion
Globe skimmer dragonflies are known to occur in Japan during
the summer, and it has been assumed that those seen early in the
season, in April and May, are arrivals from further south, whereas
Fig. 5. Distribution through time of wing δ 2H values measured for Pantala
flavescens collected at two sites in Japan in 2017: Kagoshima 9–30 April,
Kagoshima 30 May–9 June, Shizuoka 24 July–5 August, Kagoshima 5
October–12 November, and Shizuoka 24 October–13 November. Also
indicated is the mean and SD of the expected value for this species if
produced in Japan.
Table 2. Body mass (g) showing significant statistical (Wilcoxon rank-sum test) differences between expected Japanese origin and immi-
grant for Pantala flavescens captured in 2017
Site Date
Kagoshima 9–30 Apr.
Shizuoka 24 July–5 Aug.
Shizuoka 24 Oct.–13 Nov
All specimens
5
dragonflies seen later in the season are locally produced (Sugiura
et al. 1999; Arai 2007, 2012; Iwata et al. 2009). Our study con-
firms once again the general phenomenon of long-distance migra-
tion in P. flavescens and provides evidence that individuals arriving
in Japan during the spring are indeed immigrating from south of
Japan. However, our investigation does not support the idea that
most of the individuals seen in Japan during the rest of the summer
are locally produced, instead suggesting that the majority are also
immigrants.
Our spring specimens, from Kagoshima in April 2017, showed
the highest δ 2Hw of any of our samples. Although this was theoret-
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ically consistent with many of these dragonflies having originated
in the main islands of Japan (Figs. 5 and 6a), this cannot have been
the case, since (as noted above) it is too cold for P. flavescens to
overwinter in Japan, except the far southern islands (where num-
bers seem too small to account for the repopulation of the rest of
Japan). Other potential areas of origin (the Korean peninsula, north
and central China, Mongolia, northern India, and the Himalaya) are
likewise excluded because they are too cold and, in some cases, also
too dry for P. flavescens reproduction in the preceding weeks and
months. That leaves two potential sources for our April sample, both
well south of Japan: south China and the northern part of Southeast
Asia (north Myanmar, Laos, and Vietnam) or the great islands of
Borneo and Sulawesi. Wind trajectory analysis (Supp Fig. 1 [online
only]; see also Tanoue et al. 2013) and studies of other insects (Mills
et al. 1996) suggest that the former is the most likely, although the
second option cannot be entirely excluded. This finding is also con-
sistent with observations from northern China (on Beihung Island
in 38°N, at the same latitude as Miyagi Prefecture), which found a
net northward migration of P. flavescens in early summer (Feng et al.
2006, Cao et al. 2018).
During early summer, in June and July, our samples showed
δ 2Hw values consistent with most individuals having originated
either from north China and Mongolia or from south China and as
far west as the Tibetan plateau (Fig. 6b and c). This is the time of the
onset of the East Asian summer monsoon and the annual Baiu rains
(Wang and Lin 2002, Yihui and Chan 2005, Sampe and Xie 2010,
Guan et al. 2020). The rains are associated with strong SW winds
above about 1,000 m (e.g., Sampe and Xie 2010: Fig. 15; Guan
et al. 2020: Fig. 1), which are known to carry insects from southern
China (Seino et al. 1987, Watanabe and Seino 1991). These same
winds provide a means for P. flavescens to reach Japan from cen-
tral south China and as far west as the Tibetan plateau. Indeed, the
frequent arrival of P. flavescens in June has previously been linked
with westerly winds associated with the Baiu rains (Asahina and
Turuoka 1967).
Our samples from late summer, during August and September
(Figs. 3 and 4), again show little evidence that they included local,
Japanese-origin dragonflies, instead showing strong evidence of
Japanese origin Immigrant P-value
0.1597 ± 0.042 0.1977 ± 0.043 0.013
(n = 23) (n = 11)
0.1414 ± 0.027 0.2441 ± 0.034 0.004
(n = 4) (n = 8)
0.1486 ± 0.026 0.1916 ± 0.04 0.012
(n = 6) (n = 34)
0.1555 ± 0.038 0.1908 ± 0.045 <0.001
(n = 33) (n = 99)
6 Environmental Entomology, 2020, Vol. XX, No. XX

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Fig. 6. Probable origins of Pantala flavescens collected in 2017 at (a) Kagoshima prefecture 9–30 April, (b) Kagoshima 30 May–9 June, (c) Shizuoka 24 July–5
August, (d) Kagoshima 5 October–12 November, and (e) Shizuoka 24 October–13 November. The legend indicates the number of individuals assigned to a given
pixel consistent with the 2:1 odds criterion chosen (see Materials and Methods).

immigration from the west. This is not surprising as the prevail- the peak of the typhoon season. From an ocean weather ship located
ing westerly winds of early summer continue into later summer. in the Pacific Ocean (at 29°N, 135°E), P. flavescens were observed
However, a particular feature of August and September is that this is regularly during summer south-westerly winds (Hashimoto and
Environmental Entomology, 2020, Vol. XX, No. XX
Asahina 1969), although particularly large numbers occurred during
late summer when a typhoon passed (Asahina and Turuoka 1968).
On mainland Japan, the association of globe skimmer dragonflies
with typhoons has long been recognized (Wakana 1959), whereas
more recently, the occurrence of other insect species with typhoons
has also been reported (e.g., Murata et al. 1998, Kohno et al. 2004).
During the course of our study, a total of six typhoons struck Japan
during August and September 2016, the second highest number
of typhoons to have hit Japan in a single year from 1951 to 2019
(Japan Meteorological Agency 2020). Our isotopic results are con-
sistent with the P. flavescens specimens collected in Miyagi, Shiga,
Saitama, and Hokkaido prefectures in August–September 2016, all
having been carried by typhoons from regions to the west of Japan.
Data from our October and November samples (Figs. 5 and 6d
and e) suggest that several individuals may have been of Japanese
origin, which does lend some support to the findings of previous
studies. For example, Tanaka and Higashi (2003) reported that
P. flavescens individuals encountered in October in Kumamoto
Prefecture (adjacent to Kagoshima) represented the locally pro-
duced third generation of the year. Nevertheless, our data also
suggest that many individuals collected in October and November
originated outside of Japan (Figs. 5 and 6d and e). At Beihung
Island (in northern China at 38°N), net migration of P. flavescens
at the end of the summer is to the southeast, toward south central
China, not to the west toward Japan (Feng et al. 2006, Cao et al.
2018). This perhaps suggests that any immigrants to Japan at this
time may have arrived from further west or north of Beihung, pos-
sibly from the Korean peninsula, northern China, the Russian Far
East, or Mongolia, a suggestion supported by our trajectory ana-
lysis (Supp Figs. S2 and S3 [online only]).
Body mass of immigrant dragonflies was significantly higher
than that of individuals of possible Japanese natal origins. These re-
sults are consistent with the findings of Anholt et al. (1991) who re-
ported that body mass in migratory Anisoptera is significantly higher
than in non-migratory forms. The same applies to the migratory
green darner dragonfly Anax junius Drury (Odonata: Aeshnidae)
of North America, in which fall migrants have a higher fat content
than local breeders (May and Mathews 2008, May 2013). Our find-
ings are also consistent with those of Ishizawa (2007) who found
that P. flavescens specimens from Saitama Prefecture were heavier
in July and August (when samples may have included mostly ma-
ture immigrants) than in September and October (when samples
may have been dominated by immature, locally bred individuals).
In addition, individuals collected during our study from Kagoshima
Prefecture in April 2017 which had lower δ 2Hw values (suggesting
that they had traveled furthest to Japan) tended to be heavier than
those with higher δ 2Hw values, and the latter individuals were also
more like to be immature (Supp Fig. S4a [online only]). This might
suggest that immature individuals, which were still gaining weight,
were locally produced. However, even the closest source of immi-
grants in April would be the Okinawa islands, some hundreds of
kilometer to the south, which suggests that none of the P. flavescens
in this sample could be considered to be genuinely locally produced.
Furthermore, our October–November 2017 sample from Shizuoka
Prefecture does not show such a relationship (Fig. 4b). We also
note that Ishizawa (2007) reported that females were consistently
heavier than males, in contrast to our findings. In short, this consid-
eration of body mass and maturity may be a promising approach
for future studies involving more and larger samples, but it has not
provided unequivocal insights in this case.
The use of δ 2H analyses of insect wings in combination with
known precipitation δ 2H isoscapes to infer natal origins of
7
migratory species is becoming an established approach to inves-
tigating the ecology of migratory insects in general (reviewed by
Hobson et al. 2018, Hallworth et al. 2018) and that of P. fla-
vescens in particular (Hobson et al. 2012a, Cao et al. 2018). Using
a single isotope to infer origins at continental scales, however, ne-
cessarily can be ambiguous and is entirely dependent on the pat-
tern of the underlying precipitation δ 2H isoscapes. Nonetheless,
the strong isotopic signal associated with the Himalayas, the
Tibetan Plateau and northern China is particularly useful, and
the phenology of emergence of cohorts from those regions de-
serves closer inspection as they may drive the demography of the
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species across the wider region, including Japan (cf., Anderson
2009, Hobson et al. 2012a). Other isotopes to consider in future
include those of strontium, carbon, and nitrogen, but this would
involve the derivation of predictive isoscapes for these elements
or the establishment of thresholds related to land-use practices
(Wassenaar 2019). For example, dragonflies would be expected to
be enriched in 15N if they were exposed to substantial agricultural
influences, while strontium isotopes are related to, often complex,
geological history.
To date, studies assigning individual P. flavescens to origins
using δ 2Hw analyses have used calibration algorithms linking δ 2Hw
with the expected amount-weighted mean annual precipitation δ 2H
(Hobson et al. 2012a, Cao et al. 2018), and there is generally good
agreement (i.e., r2 > 0.7). However, P. flavescens has a relatively
short larval period (of only about 6 wk) compared with the species
that were used in the calibration exercise of Hobson et al. (2012b).
A more appropriate species-specific calibration algorithm for P. fla-
vescens should be developed. Further refinements to wing δ 2H isos-
capes for P. flavescens could use GIS layers to link more realistic
origins seasonally with various population cohorts, winds, and other
environmental factors. For example, a spatially explicit isotopic sur-
face related to the extent of rice agriculture in east and southeast
Asia might better inform origins of dragonfly populations there (Cao
et al. 2018) and ultimately contribute to our understanding of life
history and population dynamics of P. flavescens.
Supplementary Data
Supplementary data are available at Environmental Entomology
online.
Acknowledgments
Blanca X. Mora Alvarez assisted with sample preparations for stable isotope
analyses that were conducted at the LSIS-AFAR stable isotope laboratory of
the University of Western Ontario. We also acknowledge Noriaki Nunokawa,
Naoya Ishizawa, Akiko Minagawa, Motoharu Fukui, Mikito Okazaki, Hisao
Imamura, Kunihiko Matsuhira, Taku Tsukada, Kouji Nakamine, Hidetoshi
Sugita, Kazunari Futamachi, and Kazuya Yoshida of their assistance in col-
lecting samples of Pantala flavescens.
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