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Aquatic Mammals 2015, 41(3), 327-332, DOI 10.1578/AM.41.3.2015.327
Short Note
Insights from a Gray Whale (Eschrichtius robustus)
Bycaught in the Taiwan Strait off China in 2011
Wang Xianyan,
1
Xu Min,
1, 2
Wu Fuxing,
1
David W. Weller,
3
Miao Xing,
1
Aimee R. Lang,
3
and Zhu Qian
2
1
Third Institute of Oceanography, State Oceanic Administration, Xiamen 361005, China
2
Ocean College, Shandong University at Weihai, Weihai 264209, China
E-mail: qianzhu@sdu.edu.cn
3
Southwest Fisheries Science Center, National Oceanic and Atmospheric Administration, La Jolla, CA 92037-1022, USA
Wang Xianyan and Xu Min equally contributed in this work.
Gray whales (Eschrichtius robustus) are recog-
nized as two distinct populations in the North
Pacific Ocean: the eastern North Pacific (ENP)
population (also known as the California-Chukchi)
and the western North Pacific (WNP) population
(also known as the Korean-Okhotsk) (Andrews,
1914; Rice & Wolman, 1971; LeDuc et al., 2002;
Weller et al., 2013b). Gray whales in the Atlantic
were extirpated prior to the 19th century (Mead
& Mitchell, 1984). The ENP population migrates
from winter areas off Baja California, Mexico, to
summer feeding areas in the Bering, Beaufort, and
Chukchi Seas (Swartz et al., 2006; Weller et al.,
2013b). This population was removed from the
U.S. List of Endangered and Threatened Wildlife
in 1994 and is currently estimated to number
approximately 21,000 individuals (Durban et al.,
2013). The WNP population feeds in the Okhotsk
Sea off Sakhalin Island, Russia (Weller et al.,
1999, 2002), and also off the southeastern coast of
the Kamchatka Peninsula (Tyurneva et al., 2010).
The wintering grounds of the WNP population
remain unconfirmed but may be along the coast
of southern China based on sighting, stranding,
and catch records (Wang, 1984; Omura, 1988;
Henderson, 1990; Kato & Kasuya, 2002; Zhu,
2002; Weller et al., 2013a). A recent population
assessment using an individual-based stage-struc-
tured model resulted in a median estimate of 140
± 6 (SE) 1+ (non-calf) individuals for the WNP
population in 2012 (Cooke et al., 2013). This
population is redlisted by the International Union
for Conservation of Nature (IUCN) as Critically
Endangered (Reilly et al., 2008).
During the winter, some gray whales seen feed-
ing off Sakhalin and Kamchatka have been observed
to migrate to the west coast of North America,
including Mexico (Weller et al., 2012; Urbán
et al., 2013), while others migrate to portions of
the WNP off Asia, including the coasts of Japan
and China (Weller et al., 2008, 2013a). Despite
this level of mixing, genetic comparisons of ENP
and WNP gray whales have found significant dif-
ferences in both nuclear and mitochondrial DNA
(LeDuc et al., 2002; Lang et al., 2010, 2011) sup-
porting their management as distinct population
stocks (Weller et al., 2013b).
Gray whales are exceptionally rare off China,
with only 24 sighting, stranding, or capture records
since 1933 (Weller et al., 2013a). The gray whale
was listed in China as Category II of the National
Key Protected Animals in 1988 and was listed
under the Chinese Red List of Endangered and
Threatened Wildlife and Plants in 1994. The most
recent record of a gray whale from China is from
5 November 2011 when a dead female gray whale
(Figure 1) was found entangled in a set gillnet off
Fujian Province in the Taiwan Strait (Figure 2).
This specimen was 13.1 m in length and 21,000 kg
in weight, and represents the first gray whale
record from Chinese waters in the 21st century. To
explore the possibility of linking the 2011 Taiwan
Strait gray whale to other areas prior to its death,
skin tissue of the whale was collected for genetic
analysis, and photographs of the whale were com-
pared to the existing gray whale photo-identifica-
tion catalogs for the WNP and ENP.
Total DNA was extracted from skin tissue of
this 2011 Taiwan Strait specimen following the
recommended protocol of the TIANamp Genomic
DNA Kit (Catalog No. DP304, TIANGEN). The
extracted DNA was stored in TE buffer (10 mM
Tri-HCl, 1 mM EDTA, pH8.0). The mtDNA control
region was amplified through the polymerase chain
328 Wang et al.
reaction (PCR) using the following primer set: P1:
5’-GAATTCCCCGGTCTTGTAAACC-3’ and P2:
5’-TCTCGAGATTTTCAGTGTCTTGCTTT-3
(Hoelzel et al., 1991). PCR was carried out in a
total volume of 25 μl comprising 100 to 400 ng of
DNA template, 0.2 mM of each dNTP, 0.4 μM of
each primer, and 1.25 unit of ExTaq polymerase
(TaKaRa). The amplification program was con-
ducted as follows: preheating at 94° C for 5 min;
cycling 38 times at 94° C for 40 s, 50° C for 30 s, and
72° C for 1 min; and extending at 72° C for 5 min.
The PCR products were electrophoresed
through a 1% Agarose gel to confirm successful
amplification, and then the PCR products were
sent to Shanghai Invitrogen Biotechnology Co.,
Ltd. for direct sequencing. Cycle sequencing was
performed using the above primers, and then both
strands were sequenced independently on an ABI
3730 Automated DNA Analyzer. The resulting
sequences were edited and spliced by BioEdit,
Version 7.2.5 (Hall, 1999) with manual correction.
PCR amplification and sequencing were repeated
separately again to account for lab error. Finally,
a complete control region (D-loop) sequence, 934
bp in length, was obtained. The full length of the
D-loop sequence was used for the BLAST search
(http://blast.ncbi.nlm.nih.gov/Blast.cgi). BLAST
results indicated that the 522 bp sequence repre-
senting the 5’ end of the control region was com-
pletely identical to the haplotype R (Figure 3)
described in LeDuc et al. (2002) and reported as
haplotype 18 in Lang et al. (2011). Comparison
of the full-length sequence generated from the
China specimen with the three available gray
whale sequences (AP006471.1, AJ554053.1, and
X72200.1) that included the complete control
region indicated that the majority of variable sites
were found within the 522 bp 5’ region and, thus,
did not yield further insight into the origin of the
2011 Taiwan Strait specimen.
To date, mtDNA haplotype data are available
for 377 gray whales considered part of the ENP
population and 142 gray whales sampled in the
WNP (LeDuc et al., 2002; Lang et al., 2011).
The haplotype R was found in 12 ENP individu-
als, the majority (n = 10) of which were collected
from stranded or hunted individuals prior to 2011
(Lang, pers. comm.). However, haplotype R was
not identified among the gray whales sampled
in the WNP, all of which were biopsied on the
Figure 1. The left side (A) and right side (B) of the female gray whale (Eschrichtius robustus) found bycaught in the Taiwan
Strait, Fujian Province of China, on 5 November 2011
Gray Whale Bycaught in the Taiwan Strait 329
Sakhalin Island feeding ground between 1995 and
2007. This sample set included samples from 69%
of the individuals identified in that region through
2011 (n = 205), suggesting that the probability of
haplotype R occurring in the WNP population is
relatively low.
Based on mtDNA haplotype frequency data,
the 2011 Taiwan Strait gray whale specimen has
a higher probability of originating from the ENP
rather than the WNP. However, it is also plau-
sible that haplotype R is carried by one or more
of the individuals that utilize the Sakhalin feeding
ground or other parts of the WNP but have not
been sampled. To further examine this question,
photographs of the 2011 Taiwan Strait specimen
were compared to existing gray whale photo-
identification catalogs from the WNP and ENP.
Comparisons to photo-identification catalogs col-
lected in the WNP off Sakhalin Island and south-
eastern Kamchatka as well as catalogs from the
west coast of North America (Northern California,
Washington, Oregon, and British Columbia) and
the lagoons in Mexico did not produce a photo-
graphic match. Although the photographs of the
2011 Taiwan Strait specimen were suitable for
photo-identification purposes, they were not con-
sidered to be ideal and were additionally limited
to primarily the left side of the whale because
some epidermis on the right side had already been
removed as a result of decomposition (Figure 1B).
Some catalogs from WNP and ENP are based on
the right side; therefore, it is possible that image
quality limitations contributed to the lack of a
photographic match within the WNP or ENP.
Given that the photographs of the 2011 Taiwan
Strait whale did not match any of the individu-
als photographed off Sakhalin or Kamchatka, as
well as the absence of haplotype R among gray
whales sampled in the WNP, our results could
suggest that the 2011 Taiwan Strait whale was
a vagrant from the ENP. In recent years, there
have been sightings of two gray whales in the
Atlantic Ocean—one that was sighted twice in the
Mediterranean Sea in May 2010 (Scheinin et al.,
2011) and a second individual that was sighted
in Walvis Bay, Namibia, in May 2013 (Elwen &
Gridley, 2013). These whales were thought to be
vagrant individuals, most likely from the ENP,
14
Figure 2. Map of the western North Pacific showing: (1) location of a female gray whale
found bycaught in the Taiwan Strait, Fujian Province of China on 5 November 2011
(black star); (2) location of partial skeleton of a gray whale excavated from the Quanzhou
coast, Fujian Province of China in 1958 (black circle) (Li, 1997); (3) location of fossil
specimens of two juvenile gray whales found in the Penghu Channel of Taiwan Strait,
China (black triangle) (Tsai et al., 2014). The Taiwan Strait was shown in dark gray.
Figure 2. Map of the western North Pacific (WNP) showing (1) location of a female gray whale found bycaught in the
Taiwan Strait, Fujian Province of China, on 5 November 2011 (black star); (2) location of partial skeleton of a gray whale
excavated from the Quanzhou coast, Fujian Province of China, in 1958 (black circle) (Li, 1997); and (3) location of fossil
specimens of two juvenile gray whales found in the Penghu Channel of Taiwan Strait, China (black triangle) (Tsai et al.,
2014). The Taiwan Strait is shown in dark gray.
330 Wang et al.
15
Figure 3.
D-loop
sequence obtained in the present study was completely identical to the
Figure 3. The D-loop sequence obtained in the present study was completely identical to the AF326806 (the haplotype
R sequence) submitted to the GenBank by LeDuc et al. (2002) and Lang et al. (2010, 2011). An identical sequence is
highlighted in gray.
331 Gray Whale Bycaught in the Taiwan Strait
that had wandered into the Atlantic as a result of
diminished Arctic Sea ice cover in the northern
passageways. These records suggest that environ-
mental changes resulting from climate warming
may allow gray whales to recolonize areas used
historically or to disperse into new habitats suit-
able for living (Scheinin et al., 2011). Moreover,
without needing new ice-free passages, some gray
whales observed in the WNP have been confirmed
to migrate to the ENP (Weller et al., 2012; Urbán
et al., 2013). Therefore, it is also possible that
an ENP gray whale could disperse to the Taiwan
Strait where gray whales historically occurred.
Another possibility, however, is that an addi-
tional feeding ground (or grounds) exists in the
WNP, but it has not yet been identified or geneti-
cally characterized. Based on records from the
logbooks of 19th century ship-based whalers, gray
whales appear to have had a more extensive distri-
bution in the Okhotsk Sea, while no gray whales,
and very little search effort, were recorded in the
area of northeast Sakhalin Island that is currently
considered the primary feeding ground (Reeves
et al., 2008). While scant information is currently
available to evaluate this hypothesis, the possi-
bility that such additional WNP feeding grounds
could exist and might have been used by the 2011
Taiwan Strait whale highlights the critical impor-
tance of obtaining photographic and genetic evi-
dence, such as that presented herein, from any
gray whales recorded in areas of the WNP other
than the Sakhalin feeding ground.
Interestingly, a partial skeleton of a gray whale
was excavated from the Quanzhou coast of Fujian
Province in 1958 (Li, 1997), and fossil specimens
of two juvenile gray whales have also been found
in the Penghu Channel of Taiwan Strait (Tsai
et al., 2014) (Figure 2). Further, Henderson (1990)
summarized information from the 1869 logbooks
of New Bedford whaling ships while they were
on the “Chinese whale grounds” (p. 14). These
logbooks reported gray whales being sighted in
February at nearly an identical location as the
2011 Taiwan Strait specimen. These sources, in
combination with insights from the specimen
reported herein, suggest the possibility of a past
(and perhaps current) gray whale wintering area
in or near the Taiwan Strait region of the WNP.
Acknowledgments
The authors are grateful to Yu Xingguang, Zhang
Haifeng, Chen Jian, Tao Cuihua, and Zhao Liyuan
of Third Institute of Oceanography, State Oceanic
Administration, and Tong Shenhan of the Xiamen
Institute of Aquatic and Terrestrial Product for
their generous help and assistance during the gray
whale specimen making process.
Olga Tyurneva,
John Calambokidis, and
Jorge Urbán kindly coor-
dinated photo-identification matching to their
respective catalogs. This research was supported
by grants from the National Natural Science
Foundation for Young Scholars of China (No.
41206159) and the State Oceanic Administration
of China (SOA, No. 201105011).
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