AI 2021 Panutrakul Senanan

Aquatic Invasions (2021) Volume 16, Issue 4: 750–770
CORRECTED PROOF
Research Article
Abundance of introduced Pacific whiteleg shrimp Penaeus vannamei (Boone, 1931)
along the east coast of Thailand
Suwanna Panutrakul and Wansuk Senanan*
Department of Aquatic Science, Faculty of Science, Burapha University, Bangsaen, Chonburi, Thailand
Author e-mails: panusuwanna@yahoo.com (SP), wansuk@go.buu.ac.th (WS)
*Corresponding author
Citation: Panutrakul S, Senanan W (2021)

Abundance of introduced Pacific whiteleg Abstract
shrimp Penaeus vannamei (Boone, 1931)
along the east coast of Thailand. Aquatic Pacific whiteleg shrimp Penaeus vannamei (Boone, 1931), a highly significant
Invasions 16(4): 750–770, https://doi.org/10. global commodity, has been introduced for aquaculture to many countries outside
3391/ai.2021.16.4.10 its native range, including Thailand. In Thailand, this species was introduced to
Received: 9 March 2021 replace Penaeus monodon for aquaculture in 1998, and it has represented more than
Accepted: 15 July 2021 90 percent of total cultured shrimp production since 2005. The rapid expansion of
Published: 18 October 2021 P. vannamei aquaculture has led to escapes of P. vannamei into natural waters. We
examined abundance, size and sex ratio of P. vannamei caught by commercial fishing
Handling editor: Joana Dias
gears, namely trammel, trawl and push nets, to determine the extent to which this
Thematic editor: Charles Martin
species has entered the natural ecosystems in the Gulf of Thailand. Sampling was
Copyright: © Panutrakul and Senanan performed from August 2009 to October 2010 at five locations along the east coast
This is an open access article distributed under terms of Thailand. We consistently found P. vannamei in shrimp catches in all three fishing
of the Creative Commons Attribution License
(Attribution 4.0 International - CC BY 4.0). gears across all locations. The frequency of occurrence of P. vannamei ranged from
0.13 (trawl nets) to 0.99 (push nets). The relative abundance by weight of wild-caught
OPEN ACCESS. P. vannamei ranged from 0 to 5.43%, with highest mean abundance in Chanthaburi
(0.42 ± 0.72%) and lowest mean abundance in Chonburi (0.04 ± 0.15%). All 4,041
individuals collected were sub-adults or adults (total length = 82 to 212 mm, weight =
3.2 to 71.8 g), with a 0.92:1 female to male ratio. The samples consisted of approximately
22% adults, suggesting a notable proportion of potential breeders in the wild. Our
study can serve as a baseline for monitoring programs for aquaculture escapes. Our
report is among the few studies that document substantial numbers of P. vannamei
specimens throughout the study period. Participation by fishers proved valuable for
detecting P. vannamei in the wild. Incorporating a similar sampling approach in
fisheries monitoring programs will provide a consistent means of detecting
population trends of P. vannamei in the wild.
Key words: alien species, aquaculture, invasion, Gulf of Thailand, population

establishment
Introduction
Aquaculture is one of the significant and documented pathways to alien
species introduction worldwide (Molnar et al. 2008; Tricarico et al. 2016).
Aquaculture facilitates the release of alien species in large numbers, as
farming areas are typically extensive and close to natural waterways, and
frequent movements of cultured animals between distant locations can
assist their geographic spread (De Silva et al. 2009). Although alien species
Panutrakul and Senanan (2021), Aquatic Invasions 16(4): 750–770, https://doi.org/10.3391/ai.2021.16.4.10 750
Abundance of introduced P. vannamei in Thailand
introduced for freshwater aquaculture can be extremely invasive (e.g.,

Tricarico et al. 2016), comparable data for those introduced for
mariculture are quite limited. In this study, we examine the extent to which
the Pacific whiteleg shrimp Penaeus vannamei (Boone, 1931), one of the
world’s most cultured marine species, has spread into coastal areas of
eastern Thailand during its years of peak aquaculture production.
Penaeus vannamei has become a major global commodity, with its
production exceeding 4,000,000 tonnes in 2016 (FAO 2018), and with
Southeast Asia being an important producer (1,483,935 tonnes). Aquaculture
of P. vannamei has rapidly expanded worldwide since the early 2000s, and
major producing countries, including Thailand, are outside of its native
range. This species is native to the western coast of Central and South
America, from Mexico to Peru (Holthuis 1980; Perez Farfante and Kensley
1997). In its native habitats, P. vannamei can tolerate a wide range of
salinity (i.e., euryhaline species) and temperature (Wyban and Sweeny
1991; Medina-Reyna 2001).
In Thailand, P. vannamei was introduced for commercial production in
1998 (Briggs et al. 2004) to mitigate the economic losses due to disease
outbreaks in a native shrimp species, Penaeus monodon Fabricius, 1798.
Because of its fast growth, tolerance to high rearing densities and high
disease resistance, P. vannamei has been widely cultured in both inland
facilities and along the coast of Thailand. Production of P. vannamei in
Thailand has risen rapidly from less than 1,000 tonnes in 2001 to
approximately 600,000 tonnes in 2011. Since 2005, P. vannamei production
has accounted for more than 90 percent of the total marine shrimp culture
production in Thailand. Despite recent disease outbreaks that have impacted
Thailand’s shrimp aquaculture, the annual production has remained at
300,000–400,000 tonnes since 2014 (Department of Fisheries 2019).
Rapid expansion of the aquaculture of this alien shrimp species in Thailand
during the early 2000s led us to examine possible escapes of P. vannamei
into natural waters. We first observed the presence of juveniles and sub-
adults in Thai natural waters in 2005 (Senanan et al. 2007), and subsequently
detected increased numbers and size of wild-caught individuals in the
Bangpakong Estuary in 2006 and 2007 (Chachoengsao Province, eastern
Thailand, Senanan et al. 2009; Panutrakul et al. 2010). Panutrakul et al.
(2010) and Chavanich et al. (2016) showed that P. vannamei can survive in
the environmental conditions found in Thai waters, and may compete for
food with native species.
Escaped P. vannamei have also been detected in natural ecosystems of
Texas, USA (Balboa et al. 1991), Vietnam (Luu and Thanh 2005; Binh et al.
2009), Brazil (Loebmann et al. 2010; Barbieri et al. 2016), the southern Gulf
of Mexico (Wakida-Kusunoki et al. 2011) and India (Roshith et al. 2018).
Due to habitat similarity between the recipient and native environments,
researchers suspected a high likelihood for a self-sustaining population and
have voiced concerns over a suite of potential ecological effects (e.g.,

Balboa et al. 1991; Senanan et al. 2010; Chavanich et al. 2016). However,
neither the realized impacts of escaped P. vannamei nor the establishment
of a self-sustaining population have been sufficiently documented in any of
these locations. In addition, P. monodon, another important penaeid shrimp
species for mariculture, has established populations in non-native habitats
along the coast of Nigeria (e.g., FAO 1999; Yakubu and Ansa 2007; Oketoki
2015; Komi and Francis 2016). These observations, along with reports on
increasing numbers of P. monodon in western North Atlantic and the Gulf
of Mexico (Fuller et al. 2014; Sandoval et al. 2014) and the Egyptian
Mediterranean (Khafage et al. 2019), suggest that Penaeus shrimps can
potentially be invasive.
This study expands on earlier work in the Bangpakong Estuary and
coastal areas of Chonburi and Chachoengsao provinces (< 3 km from shore,
Senanan et al. 2007; Panutrakul et al. 2010) to include the entire eastern
coast of Thailand (four provinces). During the upward production trend in
2006–2011, the eastern coast of Thailand harboured approximately 29% of
the P. vannamei farming area (17,115 hectares) in the country, and
contributed to a similar percentage of the annual production (153,118
tonnes, averaged across 2006–2010). In addition to a high likelihood of
escape by shrimp due to intense farming activities, eastern Thailand’s long
coastline offers diverse coastal and marine habitats accessible to escapees.
Our research, therefore, set out to monitor the presence of P. vannamei
along the eastern coast of Thailand in nearshore to offshore habitats. From
August 2009 to October 2010, we examined abundance, size distribution,
and sex ratio of P. vannamei in commercial harvests by trawls, push nets,
and trammel nets, covering fishing grounds in estuaries to areas more than
5 km offshore in five locations along the east coast of Thailand. Also, we
adjusted the sampling techniques and data collection to improve the
detection of P. vannamei in the wild. Participation of fishers in sample
collection was a key factor contributing to informative results. Our findings
provided insights on available offshore habitats for this species outside of
its native range, and on the abundance of adults entering a potential breeder
pool. In addition, the approach used in this study can serve as a basis for
designing an effective monitoring program.
Materials and methods

Data and sample collection
For data and sample collection, we relied on the participation of local
commercial shrimp fishers/aggregators in five locations, including one site
in Chonburi Province, two sites in Chanthaburi Province, and two sites in
Trat Province (Figure 1). We focused on shrimp fisheries and relevant fishing
gear types, including trawls, push nets and trammel nets. Each gear type
Figure 1. Map of five sampling sites examined in this study (CHN-Trawl-Trammel, CHT-Trawl-
Trammel, TRT-Trawl-Trammel, CHT-Push, and TRT-Push) and respective fishing grounds for
each gear type, namely trawl nets, trammel nets and push nets. The fishing grounds were
identified based on interviews with participating fisher groups. The buffer zone within three
kilometers from shore is reserved for artisanal fisheries based on the Fisheries Act B.E. 2490.
targets fishing grounds at different distances from shore. Motorized vessels

equipped with trawl nets target fishing grounds at least 5 km from shore,
while those equipped with push nets and trammel nets target the zone 3–5 km
from shore and within 3 km of shore, respectively.
Each sample at a sampling location consisted of shrimp landings by one
or two fishing gear types, designated by a location code (Figure 1, Table S1).
The Chonburi sampling site (CHN-Trawl-Trammel) consisted of harvests
by trawl and trammel nets. The two Chanthaburi sampling sites contained
harvests by different gear types, with the site at Pak Nam Laem Sing
subdistrict (CHT-Trawl-Trammel), containing harvests by trawl and trammel
nets, and the one at Ko Proet subdistrict (CHT-Push), caught by push nets.
The two Trat sampling sites also contained landings by different gear types,
with the site at Nong Khan Song subdistrict containing harvests by push
net (TRT-Push) and the site at Khlong Yai subdistrict containing harvests
by trawl and trammel nets (TRT-Trawl-Trammel).
During August 2009 to October 2010, data on abundance and size of
wild-caught P. vannamei were collected monthly (0–29 fishing trips/month,
depending on gear type and location, Figure 2), with the cooperation of
commercial fishers. The first group (group I) of participating fishers
specialized in trawl fisheries or trammel netting and contributed to the
samples at CHN-Trawl-Trammel, CHT-Trawl-Trammel and TRT-Trawl-
Trammel. For this group, we visited the sites once a month and examined
the harvests from the fishing trips occurring on the days of visit (0–10
fishing trips for trawls and 0–4 fishing trips for trammel netting per month).
Figure 2. Total number of fishing trips examined for each fishing gear type per month at sampling
locations.
The second group (group II) specialized in push-net fisheries and assisted
sample collection at CHT-Push and TRT-Push. In contrast to Group I, this
group continued to assist in collecting daily catch data and the specimens
in catches from fishing trips that occurred outside our monthly visits. All
specimens were kept in a freezer until our monthly visits. Once a month,
we examined the catches on the day of the visit and collected specimens
from the fishers (2–29 fishing trips per month). This assistance provided us
with a larger sample from push nets than from the trawl and trammel net
fisheries.
Penaeus vannamei were typically present in commercial shrimp catches
along with P. merguiensis, P. monodon, P. semisulcatus, Metapeneaus spp.
and other small shrimps such as Metapenaeopsis spp., Parapenaeopsis
hungerfordi and Trachypenaeus spp. Because of the morphological similarity
between wild-caught P. vannamei and banana prawn, P. merguiensis
(Figure 3), P. vannamei were often unrecognized and sorted as P. merguiensis.
Prior to the research, we trained fishers to recognize the morphological
distinction between the two species. The critical diagnostic characteristics
between the two species included (1) the length and shape of rostrum: the
rostrum of P. merguiensis is short but high and triangular in shape, while
the P. vannamei rostrum is longer and more straight; (2) the length of
antennular flagellum of P. vannamei is very short compared to that of
Figure 3. Diagnostic morphological characteristics between native P. merguiensis (top) and

introduced P. vannamei (bottom). Key diagnostic characteristics include (1) the shape of rostrum,
(2) the length of antenula flagellum, (3) the colour of walking legs (pereopods) and (4) the
colour of uropods (Perez-Farfante and Kensley 1997).
P. merguiensis; (3) the colour of walking legs (pereopods): P. merguiensis

pereopods are yellowish, while P. vannamei pereopods are reddish; and
(4) the colour of uropods: uropods of P. merguiensis are reddish, while those
of P. vannamei are more greenish (Perez-Farfante and Kensley 1997).
All of the wild-caught P. vannamei, both from our research team and the
fishers, were transported to the Department of Aquatic Science, Burapha
University, where the species of each individual was verified. For each
verified individual, we measured the total length (from tip of the rostrum
to the distal end of the telson, to the nearest 0.1 mm) and body weight (to
the nearest 0.01 gram) and determined its sex based on the presence of a
thelycum (female) or petasma (male), which are secondary sexual
characteristics. Abundance was expressed as the frequency of occurrence,
i.e., the proportion of fishing trips in which P. vannamei was found to total
fishing trips for each gear type, and as the percentage of P. vannamei
weight relative to total shrimp harvest per fishing trip (hereafter relative
abundance). We also evaluated size frequency distributions of wild-caught
P. vannamei throughout the study period in Chanthaburi and Trat, where
push net harvests provided consistent sampling and large sample sizes. To
test the effects of location and fishing gear type on the relative abundance
of P. vannamei, and the effects of these two factors and sampling period on
body size, either one-way or two-way ANOVA was performed. Tukey’s
post hoc multiple comparisons test was used to determine differences
between sample pairs.
To examine the growth pattern of wild-caught P. vannamei, the length-
weight relationship of P. vannamei was estimated using the conventional
formula: W = aLb (Kutkhun 1962), where W is the body weight in grams

(g), L is the total length in millimetres (mm), and a and b are constants.
The parameter b is also known as the allometry coefficient. It indicates the
rate of weight gain relative to growth in length, with b > 3 meaning positive
allometry (i.e., weight increases faster than length), b < 3 meaning negative
allometry (i.e., length increases faster than weight) and b = 3 meaning
isometry. We evaluated the length-weight relationship for all specimens
and for each sex separately (males: N = 2,110, total length = 100.0–200.0 mm;
females: N = 1,931, total length = 82.0–212.0 mm). To test whether the
average b value was significantly different from 3.0, a t-test was performed
at the significance level of 0.05. All statistical analyses were performed
using the statistical package IBM SPSS Statistic 19.0 (SPSS (Thailand) Co., Ltd.).
Results
We consistently observed the presence of sub-adult and adult P. vannamei
among the commercially-harvested shrimp from the eastern coast of
Thailand, with the abundance varying among locations and fishing gear
types. Our results indicate an essential prerequisite to establishing a
population of introduced P. vannamei.
Frequency of occurrence and relative abundance of P. vannamei in

commercial shrimp harvests from five locations along the eastern coast of
Thailand
We found P. vannamei in all fishing gear types at all locations, with
variation in the frequency of occurrence and the relative abundance of
P. vannamei per fishing trip among fishing gear types and sampling locations.
Native shrimp species in the harvests included P. merguiensis, P. monodon,
P. semisulcatus, Metapeneaus spp., Metapenaeopsis spp., Parapenaeopsis
hungerfordi and Trachypenaeus spp. Over the entire study period, the
frequency of occurrence of P. vannamei ranged from 0.13 to 0.99. Of the
three gear types, the highest frequency of occurrence was found in push
nets (number of fishing trips containing P. vannamei to total number of
fishing trips were 129/132 and 257/260 for CHT-Push and TRT-Push,
respectively). The trawl and trammel net harvests from the fishing grounds
in Chonburi (CHN-Trawl-Trammel) had the lowest frequency of occurrence
in comparison to harvests with the same gear in Chanthaburi (CHT-
Trawl-Trammel) and Trat (TRT-Trawl-Trammel) (Table S1). In addition,
the high level of involvement of the group II fishers (push nets) resulted in
a higher number of fishing trips examined and more consistent data
throughout the study period (Figure 2).
The relative abundance of P. vannamei in commercial shrimp harvests
ranged from 0–5.43% per fishing trip (Table S1). Total harvest was 3–494.8 kg
per fishing trip for trawl nets, 0.5 to 55.1 kg per trip for trammel nets, and
Figure 4. Mean ± SD of relative abundance (% of P. vananmei weight to total shrimp harvest

weight) in different fishing gear types and sampling locations.
0.77–193.5 kg per trip for push nets. The relative abundance significantly
differed among sampling locations and fishing gear types. Two-way
ANOVA indicated significant effects of sampling location, fishing gear
type and the interaction between the two factors on the relative abundance
of P. vannamei (p < 0.01). Tukey’s post hoc tests revealed that the mean
relative abundance of P. vannamei in commercial harvests in Chanthaburi
(all gear types) was significantly higher than Trat and Chonburi (Figure 4).
The mean relative abundance of P. vannamei in trammel nets was significantly
higher than that of other fishing gear types (p < 0.01). However, in Chonburi,
the mean relative abundance of P. vannamei in trammel nets was not
different from those caught in trawl nets.
Although push nets typically captured P. vannamei more often than
trawl and trammel nets, the mean relative abundance of P. vannamei in
trammel nets was higher than push and trawl nets. Both indices suggested
a lower abundance of P. vannamei in Chonburi than the other locations
(Table S1, Figure 4).
Size distribution, sex ratio and growth pattern of wild-caught P. vannamei

All 4,041 individuals of wild-caught P. vannamei found in this study were
sub-adults to adults, with total length and body weight ranging from 82.0
to 212.0 mm and 3.20 to 71.80 g (Table S1). Two-way ANOVA indicated
non-significant effects of sampling location and fishing gear type on
P. vannamei length and weight (p > 0.05). However, there was a significant
effect of the interaction between sampling location and fishing gear type
(p < 0.05). The P. vannamei caught by trawl nets in Chonburi and Chanthaburi
were slightly larger than by the other fishing gear types, whereas the wild
P. vannamei caught by trawl nets in Trat were smaller than those caught by
other types of fishing gear (Figure 5).
Figure 5. Mean ± SD of total length (A) and weight (B) of wild-caught P. vannamei in each fishing gear type at different locations.
Figure 6. Temporal variation of body weight of wild-caught P. vannamei during August 2009
to October 2010 collected by push nets in Chanthaburi (CHT-Push) and Trat (TRT-Push) provinces.
The sampling consistency by push nets during the entire study period
allowed an assessment of temporal variation in P. vannamei size for
Chanthaburi and Trat. The length of P. vannamei obtained from push nets
in Chanthaburi and Trat differed among sampling periods (p < 0.01)
(Figure 6). Post hoc multiple comparisons showed that the mean total
Figure 7. Frequency distribution of total length (mm) by sampling month of P. vannamei caught by push nets in Chanthaburi
(CHT-Push) and Trat (TRT-Push) provinces during August 2009 to October 2010.
length of individuals obtained from push nets in Chanthaburi between

November 2009 and February 2010 was significantly higher (p < 0.05) than
other sampling periods, and those collected between May and July 2010
were significantly lower. For Trat, the mean length of individuals caught
between December 2009 and January 2010 was significantly greater (p < 0.05),
and those obtained between March 2010 and June 2010 were significantly
lower than other sampling periods. Individuals collected by push nets in
these two areas during the dry season (October to April) tended to be larger
than those collected during the wet season (May to September). The length
frequency distribution also suggests an increase in numbers of small-size
individuals entering the fishing grounds during the wet season (Figure 7).
Figure 8. Length-weight relationships of wild-caught P. vannamei for each sex by three fishing
gear types from five locations along the east coast of Thailand.
The length-weight relationship equations of wild-caught P. vannamei

from all sampling locations from the eastern coast of Thailand were W =
5.620*10-6 *L3.058 (R2 = 0.892) for females (1,931 individuals); W = 5.570*10-6
*L3.058 (R2 = 0.885) for males (2,110 individuals); and W = 5.480*10-6 *L3.062
(R2 = 0.910) for pooled specimens. The length-weight relationship for
females did not significantly differ from that for males (p > 0.05). The
length-weight exponent (b) indicated an isometric growth pattern in wild-
caught P. vannamei (b = 3, p > 0.05) (Figure 8).
The female to male ratios of wild-caught P. vannamei slightly varied
depending on location and fishing gear type. The female to male ratios
ranged from 0.71:1 in trawl nets in Trat to 1.5:1 in trammel nets in
Chonburi and Trat, with an average female to male ratio of 0.92:1 (Table S1).
Across all locations, the female to male ratio in trawl nets (0.80:1) was
slightly lower than in push nets (0.91:1) and trammel nets (1.17:1).
Discussion
Our research demonstrates high abundance, in terms of frequency of
occurrence and relative abundance, of the shrimp species P. vannamei in
natural waters outside of its native range due to aquaculture escapes.
A noticeable proportion of adult-size shrimp in commercial catches and
the geographic spread from estuary to offshore areas also suggests an
important prerequisite to population establishment. The consistent
presence of P. vannamei in commercial catches and their increased mean
body size from earlier reports (e.g., Senanan et al. 2007; Panutrakul et al.
2010) provides strong evidence that this species can survive and reach the
adult life stage in natural environments. Higher participation of fishers
greatly improved the detection of P. vannamei in the wild.
High frequency of occurrence and relative abundance

The number of P. vannamei caught in eastern Thailand in this study (4,041
individuals) is much higher than reported in other non-native habitats
(Balboa et al. 1991; Santos and Coelho 2002; Luu and Thanh 2005; Pérez et
al. 2007; Binh et al. 2009; Junqueira et al. 2009; Wakida-Kusunoki et al.
2011; Barbieri et al. 2016; Roshith et al. 2018). These reports documented
low numbers of P. vannamei specimens (from less than ten to less than a
hundred individuals during the study periods); they rarely reported
quantitative abundance estimates. The highest numbers of P. vannamei
found previously were in Brazil, where Barbieri et al. (2016) reported 64
individuals in gerival nets in the Cananeia, Iguape and Ilha Comprida
lagoon estuary complex between March 2005 and December 2013.
In this work, we document high abundance of P. vannamei along the
eastern coast of Thailand, both in terms of frequency of occurrence
(number of fishing trips with P. vannamei/total number of fishing trips)
and relative abundance (percent P. vannamei weight in total shrimp
harvest per fishing trip). We observed variation in the frequency of
occurrence and relative abundance by fishing gear type and location. Push
netting in Chanthaburi and Trat frequently encountered P. vannamei, but
trammel netting in Chanthaburi and Trat showed higher relative abundance.
Our observations may reflect (1) differential efficiency of fishing gear
types, (2) proximity to shrimp farms and preferred habitats for sub-
adults/adults, and (3) the level of fishers’ assistance in sample collection.
Vessels equipped with push nets actively targeted shrimp stocks in
restricted fishing grounds (between 3 and 5 km from shore, due to regulations
and size of the vessels), while trawling, another active gear type, typically
covers larger and more offshore fishing grounds (> 5 km from shore, Figure 1).
Trammel netting, on the other hand, is more passive, and targets estuarine
and nearshore areas.
Higher abundance of wild-caught P. vannamei in Chanthaburi and Trat
may also reflect the species’ preferred habitats and proximity to shrimp
farms. Fishing grounds for all fishing gear types in Chantaburi and for
push netting in Trat are located at the mouths of the Chanthaburi River
and Trat River, respectively. These areas provide not only the estuarine and
marine conditions for sub-adults and adults, but also harbour extensive
mangrove habitats that could serve as nursery and feeding grounds for
escaped or naturally produced post-larvae and juveniles. In addition, these
habitats are close to high-density shrimp farming areas, located mostly in
coastal districts of these eastern provinces. During 2008–2010, Chanthaburi
and Trat had 4,730 and 2,021 hectares of shrimp farms, respectively,
compared to only 84 hectares in Chonburi (Department of Fisheries
2012a). The relative abundance of P. vannamei observed in this study
increased exponentially as the farming area (hectares) increased (R2 = 0.9694;
0,45
Chanthaburi
0,4
Relative abuandance (% by weight)

0,35
0,3
0,25
0,2
0,15
Trat
0,1
y = 0,0334 e0,0005 x
Chonburi R² = 0,9694
0,05
0
0 500 1000 1500 2000 2500 3000 3500 4000 4500 5000
Shrimp farming area (hectares)
Figure 9. Exponential relationship between relative abudance (% by weight) of P. vannamei

and shrimp farming area in provinces along the east coast of Thailand, with shrimp farming
areas in Chonburi, Chanthaburi and Trat, averaged across 2008–2010.
Figure 9). In addition, the proximity to shrimp farms may also explain the
high frequency of occurrence (0–100%) and relative abundance (5.9 ± 14.0%)
in the Bangapakong Estuary, where shrimp farming area in the watershed
(Chachoengsao and Prachinburi provinces) totalled 9,062 hectares in 2006
(Department of Fisheries 2008).
The level of assistance from fishers may have also contributed to the
detection of the large number of specimens. The fishers involved in push-
net fisheries (group II) were willing to integrate the sorting of P. vannamei
into their shrimp sorting routines (by species and size group). In addition
to the specimens caught during each visit, we also had access to specimens
encountered on other fishing trips, along with the total catch data. As a
result, we examined a large sample of fishing trips (Figure 2) and were able
to obtain consistent and adequate numbers for the temporal analysis of
different size classes (Figures 6 and 7). In contrast to group II, group I
fishers (trammel netting and trawls) only allowed us to search for
specimens on the days of our visits, and the sorting was done under time
constraints. Samples obtained from trammel netting and trawls, therefore,
likely underestimated the frequency of occurrence of P. vannamei in the
wild and induced high variability in the relative abundance estimates.
Regardless of the level of assistance, the participation of fishers
substantially improves the detection rates of the alien shrimp species in the
wild. Due to small percentages of wild-caught P. vannamei in the
commercial harvests, our approach taken earlier in the study (February to
June 2009) of sub-sampling harvests once or twice a month was not adequate.
It is not surprising that we have not seen any reports of P. vannamei in the
national fisheries statistics, as the agency’s methods rely on sub-sampling
from harvests and interviews with fishers. Furthermore, due to the lack of
economic incentives to sort out the alien species, fishers typically sell the
wild-caught P. vannamei (if any) along with other local shrimp species.
Survival and growth in natural habitats in Thailand

Salinity and temperature in the estuarine and marine ecosystems along the
eastern coast of Thailand are within the ranges of physiological tolerance of
P. vannamei at each life stage in its native habitats (Medina-Reyna 2001).
Although P. vannamei can tolerate a wide range of salinity (0.5–45 psu), its
preferred salinity is between 7–34 psu. Based on aquaculture experiments,
this species grows particularly well at low salinity of around 10–15 psu
(where the environment and the blood are isosmotic) (Wyban and Sweeney
1991). The species can also tolerate a wide range of temperature, with
optimal temperature between 23–30 °C; smaller individuals (1 g) grow best
at 30 °C, while larger individuals (12–18 g) grow best at 27 °C (Briggs et al.
2004). Typical salinity and temperature in the marine environments in the
Gulf of Thailand, a semi-enclosed part of the South China Sea, range from
23.0–33.0 psu and 26.5–32 °C, respectively (Lowwittayakorn 1988). However,
salinity and temperature in the estuarine ecosystems such as the Chanthaburi
River mouth and Trat bay (our study areas) are strongly influenced by
tropical monsoons. Typical salinity ranges are 24.5–31.20 psu and 0.13–
24.5 psu for the dry and wet seasons, respectively (Poonapa-amporn and
Thaipichitburupha 2018; Thaipichitburupha et al. 2019). The occurrence
of P. vannamei in both nearshore (in push net and trammel harvests) and
offshore areas (in trawl harvests) suggests that this species can survive both
estuarine and marine conditions for extended periods. It also may indicate
a range expansion from estuarine/nearshore areas, where the presence of
wild-caught L. vannmei may reflect recent escapes, to a more marine habitat.
Based on the size of the wild-caught P. vannamei found in this study
(total length = 82.0–212 mm), it is likely that escaped P. vannamei can
survive and grow to maturity in natural environments in Thailand. Similar
to those observed in aquaculture settings and its native habitats (e.g.,
Hutchins et al. 1979; Araneda et al. 2008; Medina-Reyna 2001), the growth
patterns of P. vannamei in the coastal areas of Thailand vary by life stage.
Sub-adults and adults tend to exhibit isometric growth (this study, b = 3.073)
and juveniles tend to have a negative allometric pattern (Bangpakong
Estuary, b = 2.89). This growth pattern is comparable to that of the post-
larvae to sub-adult life stages of P. vannamei in aquaculture ponds near
Corpus, Texas (b = 3.037; Hutchins et al. 1979) and in low-salinity ponds
at medium density (e.g., 130 individuals per m2; b = 2.99; Araneda et al. 2008).
However, the growth patterns of sub-adult/adult P. vannamei may vary
among locations, even in their native habitats. The growth pattern of adult
P. vannamei is positive allometry (i.e., increase in weight rather than length)
in natural habitats on the east coast of the Gulf of California (b = 3.155 for
females, 3.303 for males and 3.201 for pooled samples) (Rábago-Quiroz et
al. 2019) and the Gulf of Tehuantepec, Mexico (b = 3.22 for pooled samples)
(Ramos-Cruz 2011). These habitats are highly productive due to seasonal
up-welling events.
Possible migration of wild-caught P. vannamei

The Gulf of Thailand may accommodate a complete life cycle of P. vannamei,
which typically entails movements between estuaries/lagoons to marine
environments for growth and reproduction. The complete life cycle of
P. vannamei in its native habitat takes approximately 15 to 18 months
(Cervantes-Hernández et al. 2008). Post-larvae migrate from the marine
environment to coastal estuaries, lagoons or mangrove areas for protection,
feeding and growth until they reach sub-adult stage at about 3–5 months,
or a total length of 80–100 mm. The shrimp then leave the lagoon system
to return to the marine environment as adults to mature and spawn.
Migration of juveniles from lagoon/estuary to the coastal/marine system
occurs year-round, with a peak in the rainy season (June–July) in Mexico
(Gracia et al. 1997; Ramos-Cruz 2000; Medina-Reyna 2001; Cervantes-
Hernández et al. 2008, 2012, 2017).
The fishing grounds in eastern Thailand examined in the current study
may provide seasonal estuarine and marine habitats. On one hand, we
found mostly sub-adults and adults in commercial catches (total length =
82.0–212 mm); the size of these individuals are comparable to those
collected in the native coastal/marine habitats in Mexico (total length =
80– 220 mm). On the other hand, the wild-caught P. vannamei from the
Bangpakong Estuary were smaller than in this study (total length = 55.0–
172.0 mm; weight = 1.2–36.67 g; Panutrakul et al. 2010). However, the
mean size of wild-caught P. vannamei from the Bangpakong Estuary was
slightly greater than for a typical juvenile P. vannamei found in the native
lagoon habitats (e.g., Mar Muerto Lagoon, Southern Mexico; total length =
11–145 mm; weight = 0.1–26.3 g; Ramos-Cruz 2000; Medina-Reyna 2001).
This observation may be due to escapes of larger individuals from farms
into the Bangpakong River.
We also observed size variation of wild-caught P. vannamei during the
sampling period in push-net harvests (CHT-push and TRT-Push). Overall,
the average size of individuals collected during the dry season (November
to April) tended to be larger than for those collected during the wet season
(May to July) (Figures 5 and 6). The length frequency distribution (Figure 7)
suggests occasional pulses of small-sized individuals entering the fishing
grounds during the wet season, while larger individuals become less abundant.
This observation may reflect a migration pattern of juveniles/sub-adults
from estuaries to marine habitats. The timing of the movements of these
size groups is consistent with the movement pattern observed in juvenile
P. vannamei in its native habitats (Ramos-Cruz 2000; Medina-Reyna 2001;
Cervantes-Hernández et al. 2008, 2012, 2017). In addition, the juvenile/

sub-adult movements in the coastal areas along eastern Thailand also
coincided with a large volume of freshwater run-off from inland rivers and
flooding. Large volumes of freshwater could have transported these smaller
individuals from nearshore areas and lowered salinity within the fishing
grounds; larger individuals may have migrated offshore for more marine
habitats. Furthermore, unintentional releases of farmed P. vannamei by
flooding could have also contributed to these observed increases of smaller
individuals.
Possible natural reproduction and population establishment

Size and sex ratio of wild-caught P. vannamei in this study are indicative of
potentially available breeders in the natural waters of Thailand. A notable
fraction of individuals in this study weighed more than 30 g (21.9%), a
typical size for breeders used in aquaculture and also reported for wild
populations (e.g., Chow and Sandifer 1991; Calderón-Pérez et al. 2007;
Alfaro-Montoya 2010). The sex ratios reported in our study (female/male =
0.91:1 to 1.20:1) are within the range observed for natural reproduction in
native habitats of P. vannamei (2:1 to 1:2.8 in Sinaloa, Mexico, Calderón-
Pérez et al. 2007; 2.6:1 to 1.2:1 in Puerto Chiapas, Mexico, Ramos-Cruz
2009; 0.91:1 in the Gulf of Guayaquil and Esmeraldas, Ecuador, Nicolaides
et al. 2014).We noted that the sex ratio slightly differed by location, with
the ratio less than 1:1 for specimens collected in the nearshore fishing
grounds (< 3 km from shore), but with a ratio near or higher than 1:1 for
specimens collected at the offshore locations (> 3 km from shore).
Given the high intensity of shrimp farming in eastern Thailand, even
occasional escapes of farmed P. vannamei could result in a high propagule
pressure. A typical density for a shrimp pond is 70 post-larvae per m2 and
typical production is 1 kg per m2 (10,000 kg/hectares, translating to 500,000
20-g individuals) (SEAFDEC 2005). These escapes/releases, though only a
small fraction of the total farmed shrimp, may effectively sustain a wild
population. Our observations over more than a five-year period (2006–
2010, Senanan et al. 2007; Panutrakul et al. 2010; and this study) suggest
the expansion of the distribution range of P. vannamei in portions of the
Gulf of Thailand. The relative abundance of P. vannamei in commercial
catches in this study is noteworthy; it is comparable to some rarer local
penaeid species, such as green tiger prawn (P. semisulcatus) in commercial
harvests using similar gear types. The 2009–2010 Thailand fisheries statistics
reported that P. semisulcatus contributed to 8.06% and 1.55% by weight of
penaeid shrimp catches in trawl and push nets in eastern Thailand (fisheries
zone 1) (Department of Fisheries 2012b). In addition, the catches in trammel
nets reported for the entire Gulf of Thailand consisted of approximately 0–
0.2% of P. semisulcatus relative to the total penaeid shrimp harvest (0–8
tonnes out of 3,700–3,900 tonnes).
Despite the persistence of P. vannamei in commercial catches, the

average harvest of P. vannamei in our samples (6.5 kg/month or 0.22 kg/day)
was still much lower than that observed in a naturalized population of
P. monodon in the Andori River System, Nigeria Delta region of Nigeria.
Komi and Francis (2016) reported P. monodon catches of 126–213 kg/day
(daily catch per unit of effort = 2.88–4.88 kg/man/hour) from March 2012
to February 2013. Our results may indicate an early phase of naturalization;
however, it is still difficult to conclude that P. vannamei can establish a
population in eastern Thailand.
To tease apart the contribution of recent farm escapes from a possible
self-sustaining population, a combination of ecological surveys in appropriate
habitats and genetic identification may be insightful. In recent years, there
has been a significant reduction in aquaculture production of P. vannamei
in Thailand due to disease outbreaks (Department of Fisheries 2019). If
future sampling detects an increased proportion of P. vannamei adults and
sub-adults in commercial catches and the presence of post-larvae in nearby
nursing grounds, it would indicate an established population. Furthermore,
because the sources (imported and domestic) of P. vannamei broodstock
used in Thailand have evolved since the species introduction, it may be possible
to detect genetic differences between feral populations and newly escaped
individuals based on hypervariable genetic markers (e.g., Ren et al. 2018).
Maintenance of a population by either natural reproduction or continuous
escapes can lead to undesirable ecological consequences. Based on laboratory
experiments, Panutrakul et al. (2010) and Chavanich et al. (2016) suggested
potential advantage of P. vannamei over local shrimp species (P. monodon
and Metapenaeus spp.) in food competition due to the more active foraging
nature of P. vannamei, but with similar vulnerability to predation as the
other species (Senanan et al. unpublished data). Furthermore, P. vannamei
can transmit an alien pathogen to native species. Panutrakul et al. (2010)
observed the presence of Taura Syndrome Virus (TSV), which was
originally found in P. vannamei, in all local shrimp species examined. Despite
these potential impacts, our data did not suggest consistent relationships
between the weight of native shrimp species in the harvests (all native species
combined) and that of P. vannamei. Penaeus merguiensis dominated shrimp
harvests by trammel and push nets, while P. merguiensis and Metapenaeus
spp. were dominant in trawl harvests. The harvest weight varied greatly
among gears and locations.
The documentation of P. vannamei in the wild from our research has
contributed to the recent revision of Thailand’s alien crustacean species
list. The Office of Natural Resources and Environmental Policy and
Planning (national authority for the Convention of Biological Diversity)
listed P. vannamei, along with Cherax quadricarinatus and Procambarus
clarkii, as economically important alien species that are likely to become
invasive (Appendix II, 2018 Cabinet resolution on measures for prevention,
control and eradication of alien species, ONEP 2018). Although the regulation
mandated that relevant agencies develop appropriate management measures,
the existing reporting system for the national fisheries statistics unfortunately
does not include P. vannamei.
Conclusions
This study reports abundance estimates of escaped P. vannamei, an alien
species introduced for aquaculture, along the eastern coast of Thailand.
During our study period (August 2009–October 2010), which was during
the upward trend of P. vannamei culture in Thailand, this shrimp was
consistently caught by commercial fishing gears along the east coast of
Thailand. We recorded high frequency of occurrence, relative abundance
and number of individuals compared to prior reports in Thailand and
elsewhere. Our sampling approach contributed to these informative
outcomes. Actively maintaining fishers’ participation substantially improved
our opportunity to detect P. vannamei in the wild, especially when the
fishers incorporated P. vannamei sorting into their processing routines.
Our study also shows that the escaped P. vannamei can survive and grow
normally in natural conditions of Thai waters, from estuaries to offshore
areas. The wild-caught P. vannamei were sub-adults and adults, of which
22% had reached breeder size. These results suggest a potential range
expansion of escapees into marine habitats and increased putative breeders
in natural waters.
The wild population of P. vannamei along the east coast of Thailand
persisted at least during our study period. The maintenance of the P. vannamei
population along the east coast of Thailand could either be the result of
continuing escapes (or releases) of farmed P. vannamei or natural
reproduction. Based on existing data, however, we did not have direct
evidence for natural reproduction.
Although P. vannamei is already on Thailand’s watch-list for potentially
invasive species, additional management actions still need to take place.
Towards this end, a well-designed monitoring program will be instrumental.
A monitoring program incorporating the participation of fishers will
enhance the intensity of sampling, and thus, maximize the chance of
detection. In addition, the significant reduction of aquaculture production
of P. vannamei in Thailand in recent years due to disease outbreaks
(Department of Fisheries 2019) may allow for assessing whether the
observed level of abundance almost ten years ago can be sustained with a
lower level of replenishment from escapes.
Acknowledgements
We are grateful for the assistance provided by all participating fishers in Chonburi, Chanthaburi
and Trat provinces. In addition, we thank Associate Professor Nongnud Tangkrock-Olan for her
assistance in L. vannamei identification, Dr. Prasarn Inthachareon for making the GIS sampling
map and Mr. David J. Anderson for reviewing the English language of the manuscript. We
appreciate the comments and suggestions of anonymous reviewers; these suggestions greatly
improved the quality of the manuscript.
Funding declaration
The research was funded by the National Research Council of Thailand (2009 to 2011).
Authors’ contribution
Conceptualization and funding acquisition: SP and WS; data collection and analysis: SP;
visualization: SP and WS; and writing: WS and SP.
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Supplementary material
The following supplementary material is available for this article:
Table S1. Frequency of occurrence, relative abundance, and number of individuals of P. vannamei.
This material is available as part of online article from:
http://www.reabic.net/aquaticinvasions/2021/Supplements/AI_2021_Panutrakul_Senanan_SupplementaryMaterial.xlsx

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Aquatic Invasions (2021) Volume 16, Issue 4: 750–770

Citation: Panutrakul S, Senanan W (2021)

Key words: alien species, aquaculture, invasion, Gulf of Thailand, population

introduced for freshwater aquaculture can be extremely invasive (e.g.,

have voiced concerns over a suite of potential ecological effects (e.g.,

Materials and methods

targets fishing grounds at different distances from shore. Motorized vessels

Figure 3. Diagnostic morphological characteristics between native P. merguiensis (top) and

P. merguiensis; (3) the colour of walking legs (pereopods): P. merguiensis

formula: W = aLb (Kutkhun 1962), where W is the body weight in grams

Frequency of occurrence and relative abundance of P. vannamei in

Figure 4. Mean ± SD of relative abundance (% of P. vananmei weight to total shrimp harvest

Size distribution, sex ratio and growth pattern of wild-caught P. vannamei

length of individuals obtained from push nets in Chanthaburi between

The length-weight relationship equations of wild-caught P. vannamei

High frequency of occurrence and relative abundance

Relative abuandance (% by weight)

Shrimp farming area (hectares)

Figure 9. Exponential relationship between relative abudance (% by weight) of P. vannamei

Survival and growth in natural habitats in Thailand

Possible migration of wild-caught P. vannamei

Cervantes-Hernández et al. 2008, 2012, 2017). In addition, the juvenile/

Possible natural reproduction and population establishment

Despite the persistence of P. vannamei in commercial catches, the

Poonapa-amporn C, Thaipichitburupha P (2018) Assessment of water and sediment quality in

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