North American Journal of Aquaculture

ISSN: 1522-2055 (Print) 1548-8454 (Online) Journal homepage: http://www.tandfonline.com/loi/unaj20

Lower Lethal Temperature for Arapaima Arapaima

gigas: Potential Implications for Culture and
Establishment in Florida

Larry L. Lawson, Quenton M. Tuckett, Katelyn M. Lawson, Craig A. Watson &

Jeffrey E. Hill

To cite this article: Larry L. Lawson, Quenton M. Tuckett, Katelyn M. Lawson, Craig A. Watson
& Jeffrey E. Hill (2015) Lower Lethal Temperature for Arapaima Arapaima gigas: Potential
Implications for Culture and Establishment in Florida, North American Journal of Aquaculture,
77:4, 497-502, DOI: 10.1080/15222055.2015.1066471

To link to this article: http://dx.doi.org/10.1080/15222055.2015.1066471

Published online: 16 Sep 2015.

Submit your article to this journal

Article views: 4

View related articles

View Crossmark data

Full Terms & Conditions of access and use can be found at

http://www.tandfonline.com/action/journalInformation?journalCode=unaj20

Download by: [Universite Laval] Date: 22 September 2015, At: 02:50

 North American Journal of Aquaculture 77:497–502, 2015

C American Fisheries Society 2015
ISSN: 1522-2055 print / 1548-8454 online
DOI: 10.1080/15222055.2015.1066471
COMMUNICATION
Lower Lethal Temperature for Arapaima Arapaima gigas:
Potential Implications for Culture and Establishment
in Florida
Larry L. Lawson,* Quenton M. Tuckett, Katelyn M. Lawson, Craig A. Watson,
and Jeffrey E. Hill
Tropical Aquaculture Laboratory, Program in Fisheries and Aquatic Sciences,
School of Forest Resources and Conservation, University of Florida/Institute of Food
and Agricultural Sciences, 1408 24th Street Southeast, Ruskin, Florida 33570, USA
Downloaded by [Universite Laval] at 02:50 22 September 2015
Abstract
Arapaima Arapaima gigas is a large, predatory, obligate air-
breathing fish native to the Amazon River basin. Aquaculture pro-
ducers have identified this species as a candidate for commercial
culture in Florida. For this species, a preliminary risk assessment
using the Fish Invasiveness Screening Kit indicated a medium to
high risk of invasiveness to parts of central and southern Florida.
However, data on lower lethal temperature were not available,
which was a data gap identified by the risk assessment. Lower
lethal temperature is an important physiological characteristic,
limiting not only the culture of tropical fishes in subtropical Florida
but also the potential for establishing nonnative populations. This
study is the first known experimental test of low-temperature toler-
ance for Arapaima. Our findings suggest Arapaima will not survive
temperatures at or below 16◦ C. When compared with established
nonnative fishes in Florida, this species is extremely cold sensitive,
similar to that of Butterfly Peacock Bass Cichla ocellaris. Although
this could present challenges to producers, it might also mitigate
some concerns of regulatory staff charged with managing aquatic
invasive species.
New species development is important to commercial aqua-
culture, diversifying production and potentially increasing prof-
itability. Understanding the factors that limit the production
of new species informs culture practices and their feasibility;
however, nonnative species new to culture may face regulatory
resistance and requirements for assessment of risks associated
with escapement and establishment (Hill 2009, 2011). For the
commercial production and establishment of nonnative fishes,
tolerance to cold temperature is a fundamental aspect informing
both culture practices and risk.
*Corresponding author: lllawson187@ufl.edu
Received April 3, 2015; accepted June 15, 2015
Arapaima Arapaima gigas (Osteoglossidae, alternatively
Arapaimidae) has high potential as an aquaculture species
(Schaefer et al. 2012). The species exhibits advantageous at-
tributes such as rapid growth, air breathing, and piscivory but
will readily accept a pelleted diet. Overfishing and population
decline have resulted in restrictions on wild harvest and in-
ternational trade (Castello and Stewart 2010). The demand for
Arapaima as a food fish has led to commercial production in
parts of South America, and the potential exists for expansion
into new regions (BioTrade Facilitation Programme 2006; FAO
2012; Schaefer et al. 2012), including Florida and other warm
regions of the United States.
Arapaima are native to the Amazon basin, including trop-
ical regions of Peru, Colombia, Ecuador, Guyana, and Brazil
(Castello and Stewart 2010). In their native range, Arapaima
inhabit large, floodplain-associated rivers, floodplain lakes, and
várzea-flooded forest (Quieroz 2000). Arapaima commonly oc-
cur in slow-flowing lentic waters, but they can utilize a range of
habitats, including river channels and floodplains, for dispersal
(Quieroz 2000; Castello 2008; Arantes et al. 2013). During the
rainy season, Arapaima migrate laterally into flooded habitats
for feeding and reproduction (Castello 2008). Males build a nest
in the substrate where spawning occurs and defend the fry for up
to 6 months (Quieroz 2000; Castello 2008; Nunez et al. 2011).
Spawning has been reported in outdoor aquaculture ponds with
fingerling production ranging from 26 to 1,063 per female; how-
ever, this is probably an underestimation of their total fecundity
(Nunez et al. 2011).
Outdoor, pond-based culture has economic advantages over
indoor, tank-based culture and may be suitable and profitable
497
 498 LAWSON ET AL.
for raising Arapaima in Florida (Losordo et al. 1998; Beem
2014). Yet, outdoor pond culture may be challenging if Ara-
paima have limited cold tolerance. Winter kills of cultured trop-
ical fishes are common in Florida, and producers may have
losses despite winter protection for ponds. Conversely, if Ara-
paima have the ability to survive subtropical Florida winters,
the invasion risk of Arapaima grown in outdoor pond culture
increases. Cold temperature in subtropical Florida is consid-
ered a dominant environmental variable limiting the potential
establishment and range expansion of tropical nonnative fishes
(Shafland and Pestrak 1982; Shafland 1995; Hill 2002). In the
tropical native range of Arapaima, temperatures remain warm
year round. For example, in the Brazilian State of Amazonas,
reported seasonal water temperatures range from 27◦ C to 31◦ C
(Castello 2008). While southeastern Florida has a climate sim-
ilar to parts of the Amazon River Basin where Arapaima are
native (Kottek et al. 2006), winter temperatures in the majority
Downloaded by [Universite Laval] at 02:50 22 September 2015
of the state fall well below this range. Therefore, determining
the lower temperature tolerance of Arapaima is essential for
both aquaculture producers and natural resource managers to
make informed decisions. We filled this important data gap by
identifying the chronic lower lethal temperature for Arapaima.
We further compared these data with those for previously estab-
lished nonnative fishes in Florida to estimate a potential range of
establishment.
METHODS
Captive-reared juvenile Arapaima were imported from an
aquaculture producer in the Iquitos region of Peru and held at
the University of Florida’s Tropical Aquaculture Laboratory,
Ruskin, Florida. Arapaima were raised in indoor recirculat-
ing aquaculture systems housed inside a thermally controlled
greenhouse at about 30◦ C for approximately 6 months. Once
the fish reached a size robust enough to survive frequent han-
dling stresses, they were moved to the experimental tanks. Eight
juvenile Arapaima were randomly chosen from a population of
uniformly sized individuals and placed into individual 190-L
tanks. The experimental fish had a mean length of 40.2 cm (SD,
0.9) and mean weight of 483.5 g (SD, 37.4). These fish were
maintained at 30◦ C for 5 d until all were actively feeding on
floating extruded pellet feed. The experimental tanks were con-
nected to an external head tank chilled to 14◦ C and individual
500-W electric heaters were adjusted to maintain temperature in
each tank independently. Fecal material and uneaten feed were
siphoned from the tanks daily and water changes were made
throughout the trial to maintain pH at 7.5 and ammonia nitrogen
levels below 1 mg/L.
We used chronic lethal methodology to identify the lower
lethal temperature, following procedures used in multiple stud-
ies of cold tolerance in Florida nonnative fishes (e.g., Shafland
and Pestrak 1982; Shafland et al. 2010); this methodology
allows stepwise reacclimation over the course of the trial
(Fields et al. 1987; Beitinger and Bennett 2000; Beitinger et al.
2000). Temperature in each of the six experimental tanks was
decreased by 1◦ C per day. Shafland and Pestrak (1982), using
these methods for 14 tropical fish species, found a sample size
of six produced the true mean (SD, ±1◦ C) lower lethal tem-
perature 99% of the time. The two control tanks remained at
30◦ C for the duration of the experiment. Water temperature in
all eight tanks was measured three times daily (morning, noon,
and evening) with a factory-calibrated meter (YSI ProODO;
YSI, Yellow Springs, Ohio). The six treatment tanks reached
the daily target temperature within 1–2 h of adjustment. A tem-
perature was considered lethal, and thus recorded, after a fish
lacked opercular activity for at least 10 min and had no re-
sponse when prodded. Temperatures associated with cessation
of feeding and loss of equilibrium were also noted. General
fish health and changes in activity were recorded as observed.
Fish were fed floating pellets twice daily, and feeding activity
was recorded. Each morning prior to decreasing the temper-
ature, feed was administered once. If no feeding activity was
observed at this point, that temperature was recorded as the
cessation of feeding. Loss of equilibrium was defined as the
point when the fish was lying on its side and unable to right
itself.
To apply the lower lethal temperature data to the potential
distribution in Florida, we used ESRI ArcMap 10 (Redlands,
California) to create a thermal map for Florida by kriging inter-
polation of water temperature from 41 U.S. Geological Survey
(USGS) gauge stations throughout Florida (Figure 1). The data
were daily mean minimum water temperatures (◦ C) averaged
for the month of January from 1995 to 2015, although for most
gauges only a subset of those years was available. For compari-
son, occurrence data of Butterfly Peacock Bass Cichla ocellaris,
an established nonnative fish with slightly greater thermal tol-
erance than Arapaima, were downloaded from the USGS Non-
indigenous Aquatic Species Database (USGS 2015) and the
Florida Museum of Natural History (FLMNH 2015) and then
uploaded into ESRI ArcMap 10.
RESULTS
After 5 d of acclimation at 30◦ C, all eight fish were actively
feeding and showed no external signs of stress. On day 11 of
the 21-d experiment at a temperature of 25.1◦ C (SD, 0.08), all
tanks were treated with 19 mL of formalin for 12 h to control
Trichodina, a ciliate protozoan, which was observed on several
of the treatment fish. On day 15 at an average temperature of
21.7◦ C (SD, 0.15), three treatment fish attempted to consume
a feed pellet but spit it out before swallowing. The following
day at an average temperature of 20.8◦ C (SD, 0.33◦ C), all treat-
ment fish ceased feeding and did not attempt to eat for the
remainder of the experiment. All treatment fish began to show
signs of stress, including a thickened slime coat, lethargic re-
actions, and reduced swimming ability, after several hours at
 COMMUNICATION 499
Downloaded by [Universite Laval] at 02:50 22 September 2015

FIGURE 1. Thermal map of Florida with water temperature zones (daily mean minimum water temperature data in Celsius) averaged for the month of January
from 41 U.S. Geological Survey (USGS) gauge stations, and sites where Butterfly Peacock Bass were observed between 1984 and 2014 (USGS 2015). The two
Butterfly Peacock Bass sites on Florida’s west coast represent a population isolated to a thermal refuge deep-water canal system, whereas the westernmost inland
site is not an established overwintering population.

17.0◦ C. Two treatment fish died within 2 h at 16.0◦ C and the
remaining four treatment fish died overnight at this tempera-
ture. Control fish remained active and feeding throughout the
experiment and displayed no obvious external signs of stress or
disease.
DISCUSSION
Implications for Culture
The limited cold tolerance of Arapaima (16◦ C) makes it nec-
essary for thermal protection of outdoor ponds (e.g., plastic
 500
Downloaded by [Universite Laval] at 02:50 22 September 2015
FIGURE 2. Average chronic lower lethal temperature for 18 species of nonnative fishes currently classified as established in the state of Florida compared with
that for Arapaima. Data sources: Shafland and Pestrak (1982), Shafland et al. (2010), Schofield et al. (2010), and Schofield and Huge (2011).
covering and pumping in warm groundwater) during winter
in Florida. These practices may still be insufficient for colder
years. Preliminary trials in Florida utilizing these methods sug-
gest covered pond culture is possible but potentially only in a
region south of Tampa and Orlando; further north, indoor culture
will be required. Compared with outdoor pond culture, indoor
aquaculture requires substantial up-front infrastructure capital
and increased operating expenses, potentially causing produc-
tion costs to exceed market value of food fish (Losordo et al.
1998; Beem 2014). Arapaima has been successfully cultured
commercially in outdoor ponds, but much of the success is from
South America, including Brazil and Peru, where winter tem-
peratures are not an issue (Roubach et al. 2003; Schaefer et al.
2012). Although few attempts have been made to culture Ara-
paima in North America, the species has obvious potential due to
a high market value, efficient feed conversion rates, and a rapid
growth rate (10–15 kg/year; Oliveira et al. 2012). Ultimately,
LAWSON ET AL.
continued research will be necessary to bring a Florida-grown
product to market.
Implications for Establishment
Although Arapaima are not currently established or intro-
duced into Florida, the potential for commercial production in
Florida led to a recent biological synopsis and application of a
risk screen using the Fish Invasiveness Screening Kit v2 (Hill
2013). The results indicated an elevated risk to portions of south-
ern Florida (i.e., south of Lake Okeechobee), but the previous
study was unable to take advantage of the current data on lower
lethal temperature. As a large-bodied piscivore, there is concern
about the potential ecological impacts if Arapaima were to es-
tablish. Speculating upon the extent of these impacts is beyond
the scope of this study; however, estimating the potential es-
tablishment range of Arapaima using the data now available is
potentially valuable.
 COMMUNICATION
A comparison with established nonnative fishes in Florida
can be useful in risk assessment, and lower lethal tempera-
tures have been defined for multiple nonnative fishes (Shafland
and Pestrak 1982; Schofield et al. 2010; Shafland et al. 2010;
Schofield and Huge 2011; Figure 2). For some, there is also doc-
umentation of current distributions and range fluctuations fol-
lowing unusually cold winters that result in winter kills (Boucek
and Rehage 2014). Our findings suggest Arapaima exhibit lim-
ited cold tolerance compared with established nonnative fishes
in Florida (Figure 2). With an average lower lethal temperature
of 15.6–15.8◦ C, Butterfly Peacock Bass are only slightly more
cold tolerant than the Arapaima (Figure 2). Butterfly Peacock
Bass have a limited established distribution in southern Florida
(Figure 1), the population expanding during warm years and
contracting after cold events (Swingle 1967; Guest et al. 1980;
Shafland 1995). During cold years, their populations are primar-
ily restricted to deepwater canals and borrow pits that provide
Downloaded by [Universite Laval] at 02:50 22 September 2015
thermal refuges, but warm years allow for range expansions into
otherwise unsuitable habitats.
On the basis of January mean minimum water temperatures,
our results indicate Arapaima would be unable to survive winter
temperatures throughout most of Florida (Figure 1). Moreover,
survivable winter temperatures for Arapaima occur in the two
warmest zones of the temperature map, which correspond to
the distribution of Butterfly Peacock Bass (Figure 1). Similar
to the Butterfly Peacock Bass, Arapaima would probably be
constrained to thermal refuge habitats with potential range ex-
pansion during warm years but be even more susceptible to
winter kills and range constrictions during colder years.
ACKNOWLEDGMENTS
We thank our colleagues at the University of Florida/IFAS
Tropical Aquaculture Laboratory (TAL) for support and assis-
tance. The experimental system used to determine cold toler-
ance was donated to the TAL by the Florida Fish and Wildlife
Conservation Commission and we thank Kelly Gestring, Mur-
ray Stanford, and Paul Shafland. Funding was provided by the
University of Florida/Institute of Food and Agricultural Sci-
ences (School of Forest Resources and Conservation, College
of Agricultural and Life Sciences, and the TAL).
REFERENCES
Arantes, C. C., L. Castello, M. Certa, and A. Schilling. 2013. Environmen-
tal influences on the distribution of Arapaima in the Amazon floodplains.
Environmental Biology of Fishes 96:1257–1267.
Beem, M. 2014. Aquaculture: realities and potentials when getting started.
Southern Regional Aquaculture Center, Publication 441, Stoneville, Missis-
sippi.
Beitinger, T., and W. Bennett. 2000. Quantification of the role of acclimation
temperature in temperature tolerance of fishes. Environal Biology Fishes
58:277–288.
Beitinger, T., W. Bennett, and R. McCauley. 2000. Temperature tolerances of
North American freshwater fishes exposed to dynamic changes in tempera-
ture. Environmental Biology of Fishes 58:237–275.
501
BioTrade Facilitation Programme. 2006. Arapaima gigas market study: current
status of Arapaima global trade and perspectives on the Swiss, French and
UK markets. United Nations Conference on Trade and Development, Geneva.
Boucek, R. E., and J. S. Rehage. 2014. Climate extremes drive changes
in functional community structure. Global Change Biology 20:1821–
1831.
Castello, L. 2008. Lateral migration of Arapaima gigas in floodplains of the
Amazon. Ecology of Freshwater Fish 17:38–46.
Castello, L., and D. J. Stewart. 2010. Assessing CITES non-detriment find-
ings procedures for Arapaima in Brazil. Journal of Applied Ichthyology 26:
49–56.
FAO (Food and Agriculture Organization of the United Nations). 2012. Cultured
aquatic species information programme, Arapaima gigas. FAO, Fisheries and
Aquaculture Department, Rome.
Fields, R., S. S. Lowe, C. Kaminski, G. S. Whitt, and D. P. Philipp. 1987. Critical
and chronic thermal maxima of northern and Florida Largemouth Bass and
their reciprocal F 1 and F 2 hybrids. Transactions of the American Fisheries
Society 116:856–863.
FLMNH (Florida Museum of Natural History). 2015. On-line col-
lection database, ichthyology. FLMNH, Gainesville. Available:
http://specifyportal.flmnh.ufl.edu/fishes/. (March 2015).
Guest, W. C., B. W. Lyons, and G. Garza. 1980. Effects of temperature on
survival of peacock bass fingerlings. Proceedings of the Annual Confer-
ence Southeastern Association of Fish and Wildlife Agencies 33(1979):
620–627.
Hill, J. E. 2002. Exotic fishes in Florida. Lakeline North American Lake Man-
agement Society 22:39–43.
Hill, J. E. 2009. Risk analysis for non-native species in aquaculture. Southern
Regional Aquaculture Center, Publication 4304, Stoneville Mississippi.
Hill, J. E. 2011. Emerging issues regarding non-native species for aquaculture.
Southern Regional Aquaculture Center, Publication 4305, Stoneville Missis-
sippi.
Hill, J. E. 2013. Risk screen for Arapaima Arapaima gigas for Florida. Florida
Fish and Wildlife Conservation Commission, Final Report, Tallahassee.
Kottek, M. J., J. Grieser, C. Beck, B. Rudolf, and F. Rubel. 2006. World
map of the Köeppen–Geiger climate classification updated. Meteorlogische
Zeitschrift 15:259–263.
Losordo, T. M., M. P. Masser, and J. Rakocy. 1998. Recirculating aquaculture
tank production systems: an overview of critical considerations. Southern
Regional Aquaculture Center, Publication 451, Stoneville Mississippi.
Nunez, J., F. Chu-Koo, M. Berland, L. Arevalo, O Ribeyro, F. Duponchelle, and
J. F. Renno. 2011. Reproductive success and fry productions of the paiche or
pirarucu, Arapaima gigas (Schinz), in the region of Iquitos, Peru. Aquaculture
Research 42:815–822.
Oliveira, E. G., A. B. Pinheiro, V. Q. Oliveira, A. R. Junior, M. G. Moraes,
I. R. Rocha, R. Rocha de Sousa, and F. H. Costa. 2012. Effects of stocking
density on the performance of juvenile pirarucu (Arapaima gigas) in cages.
Aquaculture 371:96–101.
Quieroz, H. L. 2000. Natural history and conservation of pirarucu, Arapaima
gigas, at the Amazonian varzea: red giants in muddy waters. Doctoral disser-
tation. University of St. Andrews, St. Andrews, UK.
Roubach, R., E. S. Correia, S. Zaiden, R. C. Martino, and R. O. Cavalli. 2003.
Aquaculture in Brazil. World Aquaculture 34:28–35.
Schaefer, F., W. Kloas, and S. Würtz. 2012. Arapaima: candidate for intensive
freshwater culture. Global Aquaculture Advocate (November/December):
50–51.
Schofield, P. J., and D. H. Huge. 2011 Low-temperature tolerance of two non-
native fishes (Hoplosternum littorale [Hancock 1828], Cichlasoma bimacu-
latum [Linnaeus 1758]) established in south Florida USA. Florida Scientist
74:73–83.
Schofield, P. J., W. F. Loftus, R. M. Kobza, M. I. Cook, and D. H. Slone.
2010. Tolerance of nonindigenous cichlid fishes (Cichlasoma urophthalmus,
Hemichromis letourneuxi) to low temperature: laboratory and field experi-
ments in south Florida. Biological Invasions 12:2441–2457.
 502 LAWSON ET AL.
Shafland, P. L. 1995. Introduction and establishment of a successful Butter-
fly Peacock fishery in southeast Florida canals. Pages 443–451 in H. L.
Schramm Jr. and R. G. Piper, editors. Uses and effects of cultured fishes in
aquatic ecosystems. American Fisheries Society, Symposium 15, Bethesda,
Maryland.
Shafland, P. L., K. B. Gestring and M. S. Stanford. 2010. An assessment of
the Asian Swamp Eel (Monopterus albus) in Florida. Reviews in Fisheries
Science 18:25–39.
Downloaded by [Universite Laval] at 02:50 22 September 2015
Shafland, P. L., and J. M. Pestrak. 1982. Lower lethal temperatures for fourteen
non-native fishes in Florida. Environmental Biology of Fishes 7:149–156.
Swingle, H. A. 1967. Temperature tolerance of the Peacock Bass and a pond test
of its value as a piscivorous species. Proceedings of the Annual Conference
Southeastern Association of Game and Fish Commissioners 20(1966):297–
299.
USGS (U.S. Geological Survey). 2015. Nonindigenous aquatic species database.
USGS, Gainesville, Florida. Available: http://nas.er.usgs.gov/. (March 2015).