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Journal of Thermal Biology 28 (2003) 531537

Thermoregulatory behavior and critical thermal limits of the angelsh Pterophyllum scalare (Lichtenstein) (Pisces: Cichlidae)
! Estela Pereza, Fernando D!azb,*, Sonia Espinac
! Laboratorio Acuario, Departamento de Biolog!a, Facultad de Ciencias, Universidad Nacional Autonoma de M!xico (UNAM), Mexico, e D.F. 04510, Mexico b ! ! Departamento de Acuicultura, Biotecnolog!a Marina, Centro de Investigacion Cient!ca y Educacion Superior de Ensenada (CICESE), Km. 107, Carretera Tijuana-Ensenada, 22830, Ensenada, B.C., Mexico c ! Laboratorio de Ecosiolog!a, Facultad de Ciencias, Departamento de Biolog!a, Universidad Nacional Autonoma de M!xico (UNAM) e Mexico, D.F., 04510. Mexico
a

Received 14 March 2003; accepted 9 July 2003

Abstract

(1) Final temperature preferendum of juvenile (0.91.9 g) and adult (5.212.5 g) angelsh Pterophyllum scalare were determined with acute and gravitation methods. The nal preferenda were similar, independent of the method and development stage (29.031.1 C). (2) The critical thermal maxima (CTMax) for juveniles were 36.9 C, 37.6 C, 40.6 C, 40.8 C and for adults 38.4 C, 38.6 C, 41.0 C, 42.1 C. Adult angelsh CTMax was slightly higher than in juveniles (1 C; Po0:05); the endpoint of CTMax was the onset of spasms. (3) The acclimation response ratio for both stages had an interval of 0.330.44; these values are in agreement with results for subtropical and tropical shes. (4) Therefore it is recommended that angelsh cultivation should be consistent with temperatures that do not change abruptly throughout the year and temperature maximum does not exceed 30 C. r 2003 Elsevier Ltd. All rights reserved.
Keywords: Preferred temperature; Acute and gravitation methods; Critical thermal maximum; Acclimation response ratio; Pterophyllum scalare

1. Introduction Among environmental factors, temperature is important since it determines physiological responses of aquatic organisms, limits the rates of the biochemical reactions and affects distribution. Generally, sh are
*Corresponding author. CICESE, Department of Aquaculture, Marine Biotechnology, P.O. Box 434844, San Diego, CA 92143-4844, USA. Fax: +1-52-6175-05-34. E-mail address: fdiaz@cicese.mx (F. D!az).

well adapted to environmental temperatures, leading to similarities in ecological responses to temperature (Magnuson et al., 1979). To cope with environmental temperature changes, ectotherms have the capacity of behavioral thermoregulation, which includes the active selection of a thermal optimal habitat and avoidance of unfavorable ones (Reynolds, 1979). Kelsch (1996) showed evidence that sh selected temperatures to maximize the proportion of metabolism available for growth, activity, reproduction and other biological functions. Thus, the temperature that an organism

0306-4565/$ - see front matter r 2003 Elsevier Ltd. All rights reserved. doi:10.1016/S0306-4565(03)00055-X

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prefers often agrees with the thermal optima for such physiological processes as the metabolism, growth, swimming speed and reproduction (Brett, 1971; Beitinger and Fitzpatrick, 1979). Generally, this temperature coincides with the nal temperature preferendum. Thermal preference is a species-specic response, modied by age, food availability, season, pathologic conditions, water quality, light intensity and intra- an inter-specic competition (McCauley and Casselman, 1981; Giattina and Garton, 1982) Two measures of temperature preference were dened by Fry (1947), as acute and nal temperature preferenda. Acute temperature preference usually determined within 2 h after immersion, and in this case the animals in the gradient are strongly inuenced by their acclimation temperatures. The nal preferendum is only determined after the animals have had a sufcient period of time to gravitate towards a region of thermal preference (Reynolds and Casterlin, 1979; Jobling, 1981). According to Hutchison (1976) the knowledge of the critical thermal maximum (CTMax) provides a relevant ecological and physiological index. Angelsh in nature may encounter such temperatures either temporally or spatially. CTMax may occur at different temperatures in different species, but the physiological responses are the same across a diversity of taxa (Lutterschmidt and Hutchison, 1997). The CTMax is modied by both seasonal and acclimation temperatures. Such data are useful for evaluating the thermal requirements of an organism and its physiological status (Paladino et al., 1980). For these reasons, CTMax is an excellent standard index for evaluating the thermal requirements and physiology of the organisms (Paladino et al., 1980; Lutterschmidt and Hutchison, 1997). The angelsh Pterophyllum scalare (Lichtenstein) is a cichlid in great demand due to its beauty, reproductive capacity and adaptability to captivity; in consequence, the economic potential of the species is also high (Chapman et al., 1997). Tropical sh like P. scalare is a new species for aquaculture; it is used in hobbies and marketed all over the world; however, little attention has been paid to the optimum thermal conditions required for growth, nutrition and reproduction (Degani, 1993; Blom et al., 2000). In this study, we present temperature preference and the upper thermal tolerance of angelsh P. scalare juvenile, and adults, and their acclimation response ratio (ARR) for optimizing their culture.

de Ciencias, UNAM. Adult sh had remained for six generations in 60-l glass aquaria at 2871 C (P! rez-Cruz e et al., 1998). In this study the adults (5.212.5 g) belong to this sixth generation and the juveniles (0.91.9 g) to the seventh. The sh of both stages were kept at culture temperature, the adults in 60-l and the juveniles in 4-l aquaria to a density of 1 sh l1. Angelsh were fed twice a day with commercial balanced food (Sera-Vipan) and Artemia sp. given at 5% and 1% body weight to juveniles and adults, respectively. The feeding period was 2 h, and excess was siphoned later. Partial replacements of the water were done twice a week. The characteristics of the water were the following: 28 29 C, 8.18.3 C pH, 67 mg O2 l1, alkalinity of 103120 and hardness of 187 mg CaCO3 l1. The photoperiod was of 12 h D/12 h L with a gradual dawn to dusk transition of 30 min. 2.2. Preferred temperature The thermal preference for juvenile and adult angelsh was determined in a horizontal trough of 300 cm length and 20 cm diameter, with 15 segments. A Neslab (model HX 100) thermoregulator was connected at one end of the trough for cooling, and a 1000 W heater was placed at the other end for maintaining the thermal gradient which ranged from 11 C to 38 C. To eliminate the stratication in the water column, 15 air stones were placed along the gradient. A dissolved oxygen concentration of 6.111.4 mg l1 was maintained. Fig. 1 shows that this procedure allowed that a linear thermal gradient be maintained. The organisms were not fed for 24 h prior to the trials. 2.3. Acute method The acute preference temperature was measured in 120 juvenile and 120 adult angelsh acclimated for 25

2. Material and methods 2.1. Maintenance Angelsh were obtained from the stock of sh cultured in the Laboratory-Aquarium of the Facultad
Fig. 1. Linear regression of temperatures in the 15 segments of the gradient.

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days at 20 C, 24 C, 28 C and 32 C71 C. Ten organisms of similar size, individually tagged (Ruiz and Villalobos, 1991), were introduced into the gradient in the segment having the same temperature as their acclimation temperature. For each experimental condition three repetitions were done. For the determination of the nal preferendum we used the acute method described by Reynolds and Casterlin (1979), which consisted of 15 observations counting the number of sh in each segment every 10 min period. Simultaneously, the temperature was measured with digital thermometers that were distributed equidistantly along the trough. 2.4. Gravitation method Ten sh of similar weight tagged individually (Ruiz and Villalobos, 1991) were introduced into the gradient in the segment having the same temperature as their acclimation temperature. Four repetitions were done for adult and juvenile angelsh (N total=80). The observations of the sh were made from reections in a mirror oriented at a 45 angle, recorded hourly during 24 h; simultaneously, the temperature was measured with digital thermometers that were distributed equidistantly along the gradient. Prior to these recordings, the organisms were kept 2 h in the trough to reduce the stress caused by handling. During the day the light intensity was 0.4 1016 quanta seg1 cm2; during the night a red light was used, with an intensity of 0.06 1017 quanta seg1 cm2. 2.5. Tolerance temperature The CTMax of 120 juvenile and 120 adult angelsh acclimated gradually at experimental temperatures, were determined. The sh remained at 20 C, 24 C, 28 C and 3271 C for 30 days. Each sh was placed in a 1-l glass ask provided with constant aeration, then they were introduced into a 40-l aquarium provided with a 1000 W immersion heater and permanent aeration to maintain a uniform temperature. The water was maintained at the experimental temperature for 30 min to reduce the stress produced by handling and heated at a rate of 1 C min1. The stress events registered were the loss of righting response (LRR) and the onset of muscular spasms (OS) according to the criteria of Lutterschmidt and Hutchison (1997). When the sh reached this point they were returned to their acclimation temperature. The organisms were used once only and the data for the animals that did not recover after returning them to their acclimation temperature after OS were discarded. In the angelsh, we determined the ARR dened by Claussen (1977) as DCTM=DT or the change in the CTM per degree change in acclimation temperature.

Preferred temperature data were processed with the Exploratory Data Analysis (Tukey, 1977), and they were plotted as parallel boxes; a KruskallWallis test was employed for determining if preferred temperature differences (Po0:05) occurred in repetitions when we used an acute and gravitation methods. After this was conrmed, an analysis of variance of ranks (Kruskall Wallis test) was used to analyze pooled data (Zar, 1999).

3. Results In juvenile P. scalare the thermal preference by the acute method was 30.2 C and the inuence of the acclimation temperature was not signicant in all groups (P > 0:05), (Fig. 2A). The preferred temperatures of adult angelsh were dependent on acclimation temperature

Fig. 2. Relationship between preferred temperature of juvenile (A) and adult (B) angelsh P. scalare and their acclimation temperature. The arrow denotes the acute estimated nal thermal preferendum. Median795% condence interval.

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and were found to be 27.8 C, 28.8 C and 30.0 C, for acclimation temperatures of 20 C, 24 C and 28 C, respectively. In sh acclimated at 32 C the preferred temperature was 29.4 C (Fig. 2B). In juveniles exposed

to light/dark cycle of 24 h the interval of preferred temperatures was 26.729.2 C with a median value of 29.0 C (Fig. 3A) and in the adults was 28.431.2 C with a median of 30.1 C (Fig. 3B). Differences in the nal preferendum in juvenile and adult angelsh between day and night were not observed. Both stages had similar thermoregulatory behavior (P > 0:05) (Figs. 3A and B). During CTMax trials the LRR was observed rst, and later the OS, Table 1. Values of OS were considered the end point of CTMax. As the acclimation temperatures increased, CTMax also increased in both stages (Po0:05). Adult angelsh had a thermal tolerance 1.0 C higher than the juveniles (Po0:05), but the increase in tolerance as a function of rising acclimation temperature, was similar in both development stages. Values were 3.9 C and 3.7 C for juveniles and adults, respectively (P > 0:05). The acclimation temperature ratio (ARR) was 0.40 0.46 for juveniles and 0.330.44 for adults of angelsh, respectively.

4. Discussion The range of preferred temperatures for both stages of P. scalare was independent of acclimation temperature over the range of acclimation temperatures used in this research; similar results were obtained by Badenhuizen (1967) in Oreochromis mossambicus and Kelsch and Neill (1990) in blue tilapia Oreochromis aureus. Both species of cichlids had a narrow preferred temperature range regardless of acclimation temperature. Fish have a variety of temperature-preference relationships that can be categorized into three classes on the basis of whether they are positive, independent or negative functions of

Fig. 3. Thermoregulatory behavior of juvenile (A) and adult (B) angelsh P. scalare. Median795% condence interval.

Table 1 Critical thermal maximum (CTMax) of juveniles (n 120) and adults (n 120) Pterophyllum scalare acclimated at four different temperatures Stress responses Acclimation temperatures ( C) 20 Juveniles LRR OS 34.0 (32.9, 35.1) 36.9b (36.8, 37.0) 36.2a 35:5; 36:9 38.4b 37:8; 39:0
a

24 35.9 (35.5, 36.3) 37.6 (37.3, 37.9) 36.5 35:6; 37:4 38.6 38:1; 39:1

28 38.6 (38.2, 39.0) 40.6 40:2; 41:0 38.1a (37.7, 38.5) 41.0 (40.6, 41.4)
a

32 38.8 (37.8, 39.8) 40.8b 40:4; 41:2 41.2 (39.4, 43.0) 42.1b (42.0, 42.2)

Adults

LRR OS

The underline between groups indicates a similar effect of acclimation temperature (P > 0:05). LRR: loss of righting response, OS: onset of muscular spasms. Median values and condence interval (95%) in parenthesis. a Signicant difference of LRR between juvenile and adults (Po0:05). b Signicant difference of OS between juvenile and adults (Po0:05).

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the acclimation temperature (Johnson and Kelsch, 1998). Species such as cichlids that experience low annual thermal amplitude, but may be exposed to temperature uctuations on a daily or sub-seasonal basis (short cycle) exhibit temperature-preference relationships that are independent of the acclimation temperature (Johnson and Kelsch, 1998). Therefore the nal preferenda can be used as a measure of the temperature selected by angelsh as an index of the magnitude of temperatures to which species are adapted (Johnson and Kelsch, 1998). When both stages of P. scalare are placed separately in a thermal gradient during 24 h, the sh distribute in accordance to the presumed effect of the temperature on surplus power capacity (Bryan et al., 1990). According to Fraenkel and Gunn (1961) angelsh used an orthothermokinesis orientation mechanism; that is, juveniles the interval of preferred temperatures was 26.729.2 C and in the adults 28.431.2 C. Then, the angelsh will remain within a relatively narrow temperature range decreasing speed in those temperatures that maximize their available power (Kelsch and Neill, 1990; Bryan et al., 1990; Kelsch, 1996). Preferred temperature of shes are probably those that offer the greatest scope for activity (according to Fry, 1947) and therefore, the greatest amount of available power that could be channeled into adaptative functions such activity, growth reproduction and survival (Kelsch, 1996). CTMax is considered as a measure of thermal tolerance and is determined by raising the temperature progressively from the acclimation temperature until physical disorganization occurs in response to the thermal stressor (Becker and Genoway, 1979; Paladino et al., 1980; Beitinger et al., 2000). These authors also emphasize that the rate of temperature rise employed is an important factor. In this work, the rate of change employed was 1 C min1 (Lutterschmidt and Hutchison, 1997). As the temperature increased during a CTMax test, angelsh usually display a sequence of responses the LRR followed by the sudden OS. Lutterschmidt and Hutchison (1997) considered that OS is a more meaningful end point than LRR due to its precision and greater physiological relevance. We considered the end point of CTMax as the pre-death thermal point at which locomotory movements become disorganized and a sh loses the ability to escape the conditions which may ultimately lead to its death Beitinger et al. (2000). In both stages of angelsh this point was the onset of spasms; the sh exhibited high frequency of muscular movements, rigidity of the pectoral ns and a high frequency in quivering of the opercula. These responses seem to t best the denition of CTMax as the arithmetic mean of the collective thermal points at which the endpoint was reached by individuals of a random sample of sh (Hutchison,

1961). In aquacultural practices it is of fundamental importance to know the CTMax since it is a good indicator of the thermal tolerance of the angelsh and it allows the identication of the temperatures at which the rst signs of stress occur (Paladino et al., 1980). In both stages of angelsh we determined the ARR, according to Claussen (1977), which is a convenient index of the extent of thermal acclimation achieved, the value for P. scalare was 0.330.46. For Salmo trutta, Salvelinus fontinalis, Oncorhynchus gairdneri, O. apache, and O. gilae, organisms inhabiting cool streams, the calculated ARR was of 0.090.13 (Lee and Rinne, 1980). In the darters Etheastoma abellare, E. blennioides and E. caeruleum, inhabitants in cool to warm streams the calculated ARR was of 0.180.24 (Holohowskyj and Wissing, 1985). Rajaguru (2002) obtained ARR values for estuarine shes of about 0.20 in Therapon jarbua and 0.25 in Etroplus suratensis. In Poecillia sphenops, Hernandez and Buckle (1998) reported a value of 0.36. In the mosquitosh Gambusia afnis, inhabitant of warm spring streams, the calculated ARR was 0.40 (Otto, 1973). Cox (1978) and Woiwode and Adelman (1992) obtained ARR values of 0.43 and 0.46 in Morone saxatilis and Morone chrysops hybrid. For the subtropical sh Prochilodus scrofa for two-size class organisms the calculated ARR was of 0.380.47 (Barrionuevo and Fernandes, 1995). In the catsh Ictalurus punctatus, inhabitant of warm lakes and large rivers, the ARR calculated has an interval of 0.390.63 (Cheetham et al., 1976; Bennett et al., 1998; Currie et al., 1998; D!az and Buckle, 1999). The existent data for ARR in different sh species suggest that subtropical and tropical species have higher values. These results suggest that the ARR values are dependent on the existing habitat temperatures. D!az et al. (2002) reported a similar trend in several crustacean species from different habitats. This response is typical of aquatic poikilotherms, Johnson and Kelsch (1998), that temperate species experiencing gradual longterm temperature changes would have the necessary time to make metabolic changes that would result in substantial shifts in their ranges of tolerance. On the other hand, subtropical and tropical species that experience their greatest thermal extremes over a short term must have a broad range of tolerance to survive to relatively rapid temperature changes without time for acclimation to adjust their tolerance. Knowledge of the preference and the thermal tolerance of a species are important in dening appropriate culture conditions, since this allows us to know the thermal requirements of the organisms in a short time. Besides, the information concerning the lethal temperature and that concerning the species physiological optimum are important for inferring the level of survival in a particular climate space. To cultivate angelsh we recommend those geographical zones of

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536 E. P!rez et al. / Journal of Thermal Biology 28 (2003) 531537 e Claussen, D., 1977. Thermal acclimation in ambystomatid salamanders. Compounds Biochem. Physiol. 58A, 333340. Cox, D.K., 1978. Acclimation states of juvenile striped bass held in constant and uctuating temperature regimes. In: Thorp, J.H., Gibbons, J.W. (Eds.), Energy and Environmental Stress in Aquatic Systems. Technical Information Center, US Department of Energy, pp. 703713. Currie, R.J., Bennett, W.A., Beitinger, T.A., 1998. Critical thermal minima and maxima of three freshwater game-sh species acclimated to constant temperatures. Environ. Biol. Fish. 51, 187200. Degani, G., 1993. Growth and body composition of juvenile of Pterophyllum scalare (Lichtenstein) (Pisces: Cichlidae) at different densities and diets. Aquat. Fish. Manage. 24, 725730. D!az, F., Buckle, L.F., 1999. Effects of the critical thermal maximum on the preferred temperatures of Ictalurus punctatus exposed to constant and uctuating temperatures. J. Therm. Biol. 24, 155160. D!az, F., Sierra, E., Re, A.D., Rodr!guez, L., 2002. Behavioural thermoregulation and critical thermal limits of Macrobrachium acanthurus (Wiegman). J. Therm. Biol. 27, 423428. Fraenkel, G.S., Gunn, D.L., 1961. The orientation of animals. Kineses, taxes and compass reactions. Dover Publications, Inc, New York, p. 376. Fry, F.E.J., 1947. Effects of the environment on animal activity. University of Toronto Studies Biological Series 55, Publication of the Ontario Fisheries Research Laboratories 68, pp. 162. Giattina, J.R., Garton, R.R., 1982. Graphical model of thermoregulatory behavior by shes with a new measure of eurithermality. Can. J. Fish. Aquat. Sci. 39, 524528. Hernandez, R.M., Buckle, R.L.F., 1998. Critical thermal maximum and minimum of Poecilia sphenops acclimated to different temperatures. Riv. Ital. Aquacoltura 33, 6171. Holohowskyj, I., Wissing, T.E., 1985. Seasonal changes in the critical thermal maxima of faintail (Etheostoma abellare), greenside (Etheostoma blennioides), and rainbow (Etheostoma caeruleum) darters. Can. J. Zool. 63, 16291633. Hutchison, V.H., 1961. Critical thermal maxima in salamanders. Physiol. Zool. 34, 92125. Hutchison, V.H., 1976. Factors inuencing thermal tolerance of individual organisms. In: Esch, W.G., McFarland, W.R. (Eds.), Thermal Ecology II. US National Technical Information Service, Springeld, pp. 1026. Jobling, M., 1981. Temperature tolerance and the nal preferendum-rapid methods for the assessment of optimum growth temperatures. J. Fish. Biol. 19, 439455. Johnson, J.A., Kelsch, S.W., 1998. Effects of evolutionary thermal environment on temperature-preference relationships in shes. Env. Biol. Fish. 53, 447458. Kelsch, S.W., 1996. Temperature selection and performance by bluegills: evidence for selection in response to available power. Trans. Am. Fish. Soc. 125, 948955. Kellog, R.L., Gift, J.J., 1983. Relationship between optimum temperatures for growth and preferred temperatures for the young of four sh species. Trans. Am. Fish. Soc. 112, 424430. Kelsch, S.W., Neill, W.H., 1990. Temperature preference versus acclimation in shes: selection for changing metabolic optima. Trans. Am. Fish. Soc. 119, 601610.

M! xico were the temperatures are close to 30 C. Several e investigations have noted the specic relationship between preferred temperature and the optimum temperatures for the performance of many physiological functions (Jobling, 1981; McCauley and Casselman, 1981; Kellog and Gift, 1983; Giattina and Garton, 1982). The tolerance for living in high temperature intervals is characteristic of P. scalare which shares with African cichlids, that are also thermophilic, this tolerance gives these species a good potential for cultivation in tropical areas. In this sense it is recommended that before selecting places based on the regional temperatures, the behavioral responses of the endemic species should be considered (Mart!nez-Pala cios et al., 1996). Thus, the results obtained in this study are important for improvement or for establishing angelsh cultures. Based on this we recommend to choose places with high temperatures that do not exceed 30 C and do not change abruptly through the year.

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